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.     

IDANOTHEKION GEN. N., A SYNANGIATE POLLEN ORGAN WITH SACCATE POLLEN FROM THE MIDDLE PENNSYLVANIAN OF ILLINOIS

Idanothekion glandulosum gen. et sp. n. is a synangiate pollen organ represented by approximately 30 specimens contained in coal balls from the middle Pennsylvanian of Illinois. Each synangium is composed of seven to nine elongate sporangia that are fused laterally for approximately four-fifths of their length, and are radially arranged about, and fused to, a short central column; the central column is restricted to the proximal one-third of the synangium. Distal to the column the sporangia surround a hollow central area. Dehiscence occurred by means of a longitudinal slit along the mid-line of the inner face of each sporangium. The outer walls of the sporangia have a complex histology involving an external epidermis, a middle presumably glandular layer containing scattered enlarged cells, and an inner layer made up of thin-walled parenchyma. Vascular tissue is present in the central column and outer walls of the sporangia. Each sporangium has a prominent, attenuate, multicellular tip. Large numbers of saccate pollen grains similar to those found in numerous fossil and extant coniferophytes as well as some Mesozoic pter-idosperms were produced in each sporangium. Idanolhekion resembles some synangia assignable to Paleozoic members of the Marattiales; however, the new genus compares most closely with pollen organs believed to have been produced by members of the Pteridospermales. It seems most likely that Idanothekion represents the pollen organ of some member of the Lyginopteridaceae that produced pollen of a type which up to now has not been known from Paleozoic seed ferns.     

中华水韭孢子囊发育研究

采用半薄切片法,连续观察了极度濒危级(CR)植物中华水韭大小孢子囊的发育过程,以期从无性生殖的角度,为探讨其濒危原因提供直观可靠的理论根据。结果显示:(1)中华水韭的大小孢子叶相间排列,无混生孢子囊。(2)隔丝为孢子供给营养,其体积直接影响孢子的大小、产量和育性。(3)大小孢子囊都近半数败育,小孢子囊为整齐发育,大孢子囊为不整齐发育。(4)大小孢子囊均无柄,且都不存在开裂结构,只有孢子囊壁腐烂后才能散播孢子。研究认为,中华水韭的濒危与孢子囊的发育特征密切相关,孢子囊的高频率败育、没有开裂结构以及对环境的依赖,是造成中华水韭濒危的重要因素之一;通过与近缘类群孢子囊的比较,发现仅水韭孢子的散播借助外力,对生境要求较高,即验证了水韭古老的系统学地位,同时说明水韭更具有监测生境地区环境指标的能力。     

Autochory in ferns,not all spores are blown with the wind

Dispersal is a key process in plant population dynamics. In ferns, two successive vectors are needed: the sporangium catapulting mechanism, and wind or gravity. However, some rock ferns have a growth habit that suggests a kind of autochory by placing spores on the rock surface. Moreover, some ferns show modifications of the sporangial dehiscence. To determine the role of growth habit in spore dispersal, we checked the sporangial opening mechanism and explored the spatial distribution of plants on the walls. The presence of spores of Asplenium celtibericum, a rupicolous fern, in the rock surface was checked. In addition, its sporangial dehiscence, plant size and position in the wall were analysed. Spores and indehiscent sporangia were present on walls at each sampling moment. Their highest number was found close to the plants. There was a positive correlation between crack width and plant size. However, most plants occupy the upper half of the cliffs. The growth habit of A. celtibericum is instrumental to deposit the spores over the neighbouring rock surface, thus enhancing the probability of spores to find suitable crevices for germination. Furthermore, dispersal of indehiscent sporangia might promote intergametophytic mating, and the modified sporangial opening mechanism extends the dispersive period.     

ACAULANGIUM GEN. N., A FERTILE MARATTIALEAN FROM THE UPPER PENNSYLVANIAN OF ILLINOIS

Material described by Graham as Cyathotrachus bulbaceus is believed to represent a new genus that is a common constituent of Upper Pennsylvanian coal balls. The sessile synangia of Acaulangium gen. n. are borne in a row on either side of the pinnule midrib and are composed of four to six short, tapering, laterally appressed sporangia. The sporangia have extended tips which curve over the inside of the synangium distally and delimit a small open area inside the synangium. The outer facing walls of the sporangia are two to three cells thick throughout while the inner facing walls are uniseriate. During dehiscence the sporangia separate laterally and spore release results from the rupture of a row of elongate cells along the inner sporangium midline. Among species of Scolecopteris the new genus resembles S. illinoensis and S. minor var. parvifolia but differs in its sessile synangial attachment. The additional parenchyma present between sporangial cavities in the synangia of Acaulangium, and the tendency toward bilateral symmetry suggests an early stage in the evolution of a bivalve synangium such as is present in Marattia.     

湖北蕨类植物新记录

该文报道了湖北蕨类植物1个新记录属:冷蕨属(Cystopteris Bernhardi)及对应的新记录种宝兴冷蕨(Cystopteris moupinensis Franchet); 4个新记录种:台湾剑蕨(Loxogramme formosana Nakai)、扇羽阴地蕨[Botrychium lunaria (Linnaeus) Swartz]、耳状紫柄蕨[Pseudophegopteris aurita (Hooker) Ching]、巢形鳞毛蕨[Dryopteris transmorrisonense (Hayata) Hayata]; 2个新记录变种:尾头凤尾蕨(Pteris oshimensis var.paraemeiensis Ching)和西藏粉背蕨[Aleuritopteris subvillosa var. tibetica (ChingS. K. Wu) H. S. Kung]。这些蕨类新记录在湖北的发现,大大丰富了湖北的植物区系及物种多样性,同时对这些蕨类的经济价值或药用价值进行了简要分析,为其资源的进一步合理开发和利用提供了一些理论依据。     

蕨类植物孢子囊形态 I. 鳞始蕨科

孢子囊是蕨类植物的繁殖器官, 其形态在蕨类植物的分类和系统发育研究上具有重要意义。用次氯酸钠溶液处理新鲜成熟的孢子囊, 在光学显微镜下获得清晰的孢子囊图像, 系统研究了中国鳞始蕨科4属13种孢子囊的形态特征。结果表明, 鳞始蕨科孢子囊呈椭球形, 孢子囊柄由3列细胞构成, 环带类型为垂直环带。通过分析孢子囊形态数据探讨了中国鳞始蕨科属内及属间差异。结果表明, 乌蕨属(Odontosoria)、香鳞始蕨属(Osmolindsaea)、达边蕨属(Tapeinidium)和鳞始蕨属(Lindsaea)的孢子囊环带细胞数依次减少, 囊蒴体积、唇细胞数和囊壁细胞数的变化由大(多)到小(少)依次为乌蕨属、达边蕨属、香鳞始蕨属和鳞始蕨属。孢子囊属内差异最大的是阔片乌蕨(Odontosoria biflora)与乌蕨(O. chinensis)以及香鳞始蕨(Osmolindsaea odorata)与日本鳞始蕨(O. japonica); 而达边蕨属和鳞始蕨属的属内差异则很小。研究结果为揭示鳞始蕨科系统发育关系提供了形态基础, 特别是提出阔片乌蕨和乌蕨以及香鳞始蕨和日本鳞始蕨在孢子囊形态上的差异值得进一步研究。     

REASSIGNMENT OF ASPIDIUM HETEROPHYLLUM TO A NEW GENUS IN THE FAMILY M ATONI ACE AE

A fossil fern described originally by W. Fontaine as Aspidium heterophyllum has been reexamined. Based upon characteristics of the sorus, indusium, sporangium, and spores the plant belongs to the family Matoniaceae. There is no genus within the family to which the fossil can be assigned; thus, a new genus (Delosorus) is proposed. This report increases the number of plants within the Lower Cretaceous Potomac Group that belong in the Matoniaceae.     

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.     

A new group of Early Devonian plants with valvate sporangia containing sculptured permanent dyads

As part of a study to explore diversity and disparity in Early Devonian terrestrial vegetation, several hundreds of sporangia with in situ spores have been isolated from a Lochkovian locality in Shropshire. These include a small number (seven) of sporangia showing dehiscence into four valves and containing permanent sculptured dyads, belonging to the Cymbohilates horridus complex and C. cymosus, which are recorded in coeval dispersed spore assemblages. A further, previously described, mesofossil comprises an incomplete sporangium containing C. horridus that terminates a naked isotomously branching stem with stomata. The valvate sporangia are placed in a new genus, Partitatheca, containing four species, P. splendida (type), P. horrida, P. densa and P. cymosa, their names reflecting the names of the dispersed spore species and varieties. Complex ultrastructure in the walls of the dyads is similar to that in earlier dyads in the Dyadospora complex where it provides evidence for a hepatic affinity of the earliest embryophytes, but the new taxa present a combination of bryophyte and tracheophyte characters and are considered to represent a new embryophyte lineage. General discussion includes the development of dyads, more particularly their relevance to understanding the diversity in meiotic processes, and the disappearance of dyads from the dispersed spore record prior to the Middle Devonian. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 229–257.     

Fossil evidence of eupolypod ferns in the mid-Cretaceous of Myanmar

Divergence time estimates based on DNA sequence variation of extant species suggest that eupolypod ferns were diverse already in the Cretaceous; however, fossil evidence to support this assumption remains exceedingly rare. Holttumopteris burmensis gen. et sp. nov., a fertile fern foliage fragment preserved in a piece of Albian–Cenomanian Burmese amber from Myanmar, is characterized by divided fertile leaves with catadromous, free lateral veins. Sporangia possess a vertical annulus interrupted by the sporangium stalk and contain monolete spores with a lophate perine. The sporangia occur clustered in discrete sori overarched by a laterally attached, reniform indusium. Reconstruction of ancestral character states suggests affinities of Holttumopteris to the Thelypteridaceae; however, several taxonomically relevant characters are not preserved. This discovery is nevertheless important because H. burmensis represents the first unequivocal fossil of a representative of the eupolypods from the middle Cretaceous.     

BOTRYOPTERIS PINNULES WITH ABAXIAL SPORANGIA

Small sporangia borne abaxially on pinnules attached to Botryopteris foliar members are described from coal ball petrifactions of Early Pennsylvanian age. This is the first report of laminar sporangia in this genus. Sporangia are stalked and borne singly near lateral veins on Sphenopteris-like pinnules. Individual sporangia are of the leptosporangiate type, with a lateral annulus and a dehiscence zone of thin-walled cells immediately adjacent to the annulus. Spores are small, trilete, triangular in outline, typically have blunt spines covering the exine, and correspond to the dispersed spore genera Acanthotriletes, Leiotriletes, or Lophotriletes. These sporangia and their spores are unlike previously described globose Botryopteris fructifications from the Middle and Upper Pennsylvanian, but are similar to sporangia produced by modern members of the Osmundaceae.     

SULLITHECA DACTYLIFERA GEN. ET SP. N.: A NEW MEDULLOSAN POLLEN ORGAN AND ITS EVOLUTIONARY SIGNIFICANCE

The pteridosperm (Medullosaceae) pollen organ Sullitheca dactylifera gen. et sp. n. is described from middle Pennsylvanian coal balls. The proximally fused units of the obpyriform compound synangium separate and extend distally as finger-like projections. Each projection contains 4–6 vertically oriented cylindrical sporangia arranged in pairs along the radius of the unit; each unit extends from the outer cover wall toward the center. The distal portion of the compound synangium is hollow as a result of the lateral separation of the centripetally and distally directed synangial units. About 40 tubular sporangia are present in all and dehiscence occurs along a lateral slit in each sporangium. Vascular strands are disposed around the periphery of the organ in addition to a single strand paralleling each sporangium. Two- or three-cell trichomes and stomata are present on the organ surface. Pollen of the Monoletes type is present. A paired row of sporangia in Sullitheca composing a synangial unit is considered the homologue of a paired row of sporangia in the more compact and highly evolved genus, Dolerotheca.     

Germination of resting spores in Synchytrium species parasitic on cucurbitaceae

Germination of resting spores is described in 2 Synchytrium species viz. Synchytrium lagenariae Mhatre and Mundkur and S. trichosanthidis Mhatre and Mundkur, parasitic on cucurbitaceous hosts. The resting spores of both species behave as prosorus in germination giving rise to an attached superficial sorus of sporangia. Several anomalies observed in germination are briefly described. Efficacy of methods inducing germination is discussed relative to the field conditions.     

Leonophyllum tenellum nov. gen., nov. sp., an enigmatic plant from the Early Jurassic of the Mecsek Mts (Hungary)

The Early Jurassic flora of the Mecsek Mountains is diverse, with numerous representatives of ferns, seed ferns, cycadophytes, ginkgophytes, and conifers. Its (para-)autochthonous deposition, good preservation and low collection bias has permitted researchers to save delicate, small plant remains that would generally be missing from the fossil record. These plant fossils are characterized by having a “filmy” (probably unilayered) structure, thin stalks with flat leaf-like branches and ultimate irregular segments with streamlined epidermal cells having thick cell walls. Due to the absence of sporangia, the plant remains cannot be confidently assigned to any higher plant group, although they show some similarities to thalloid liverworts with raised vegetative bodies and also have some resemblance to the fern family Hymenophyllaceae. The new genus and species Leonophyllum tenellum Barbacka et Kustatscher is erected.     

AN UNUSUAL BOTRYOPTERID SPORANGIAL AGGREGATION FROM THE MIDDLE PENNSYLVANIAN OF NORTH AMERICA

A new fertile species of Botryopteris (Botryopteridaceae: Filicales) is described from four incomplete Middle Pennsylvanian specimens. Fertile pinnae of B. cratis sp. n. consist of branched frond members bearing numerous globose sporangia. Surrounding the sporangial aggregations are larger sterile frond members (0.5-1.5 mm diam). Fertile pinnae are oval in transverse section and possess an eccentrically developed cortex composed chiefly of fibers. Some frond members show the typical botryopterid xylem configuration with three protoxylem strands. Spherical sporangia are loosely aggregated on the smallest pinnae by short, broad stalks. The annulus is band-like, two cells high, and extends transversely across the lower half of the sporangium for approximately half the circumference. Spores are oval, trilete, verrucate, and covered by a thin separable layer. Sporangium morphology is like that of Botryopteris antiqua, but the spores closely resemble those of B. globosa. The new species is unlike previously described fructifications of Botryopteris in exhibiting a small pinna system which surrounds smaller pinnae bearing sporangia in an aggregation. The new form is considered to be less specialized than previously described globosoid forms because the sporangia are much less crowded. Isolated frond members, believed to belong to the new species, have a large central arm in the pinna xylem trace that resembles the Stephanian taxon B. renaultii. Small stems attached to the adaxial surface of frond members are radial, protostelic, centrarch, and have a three-zoned cortex. The inner cortical zone contains large elongate cells with distinctive layered deposits. Stems are covered with uniseriate multicellular hairs on multicellular bases. Stems compare closely with B. mucilaginosa in histological features.     

宽带蕨属（水龙骨科）的系统学研究

对喜马拉雅和横断山地区特有的水龙骨科的宽带蕨属Platygyria Ching&S.K.Wu植物进行了形态学、分类学、生态和植物地理学的观察与分析,探讨了该属植物的系统位置、亲缘关系和生态适应性,推测宽带蕨属是随着喜马拉雅山脉的隆升,由瓦韦属Lepisorus（J.Sm.）Ching网眼瓦韦群植物适应高寒生态环境演化而来的新生类群。确定宽带蕨属有5种,即川西宽带蕨P.soulieana（Christ）X.C.Zhang&Q.R.Liu、多变宽带蕨P.variabilis Ching&S.K.Wu、耳基宽带     

FURTHER OBSERVATIONS ON ZOSTEROPHYLLUM LLANOVERANUM FROM THE LOWER DEVONIAN OF SOUTH WALES

Zosterophyllum llanoveranum was first described by Croft and Lang in 1942. This account presents the anatomy of the axes. Details of the sporangium wall in the vicinity of the dehiscence line are reported and the spores are described. The plant is compared with other members of the genus Zosterophyllum and also with those Devonian plants possessing exarch protosteles and lateral sporangia, which Banks has placed in the subdivision Zosterophyllophytina.     

A study of the genus Aleuritopteris Fée in China

As a genus, Aleuritopteris was first founded by Fée upon Pteris farinosa Forsk. in 1852. The genus had been ever since, however, practically forgotten, because it did not receive a general recognition among the fern students, who considered it either as Cheilanthes (Baker. 1897, Diels, 1900, C. Christensen 1905), or placed it in Pellaea (Hooker. 1858, Hope et Wright 1903). Ching (1941) first reinstated the status of Aleuritopteris and later followed by Copeland (1947), Panigrahi (1961, 1962.) and Pichi-Sermolli (1975). In recent years, however, some pteridologists (Nayar, 1962, K. Iwatsuki and others) are still against it as a distinct genus. Our recent study of the rich material on hand has shown Aleuritopteris to be a distinct genus, according to both the morphological characteristics and geographical distribution. We further propose the possibly evolutionary relationships of Aleuritopteris with other related genera of the Cheilanthoid ferns, as indicated by the above scheme. The genus is now represcented by a little more than 30 species, of which 25 areknown in China, among which 8 species and 3 varieties are diescribed for the first time.