DIMENSIONAL CORRELATIONS IN DEVELOPING SELAGINELLA SPORANGIA

With length of sporangia as a developmental index, the growth relationships of sporangia during differentiation were studied in strobili of Selaginella bigelovii. The strobili usually contain two rows of megasporangia and two rows of microsporangia with a mega- opposite a microsporangium at each node. Prior to the sporocyte stage a sporangium in a megasporangiate row is larger and elongates more rapidly than a sporangium opposite it at the same node in a microsporangiate row. The number of sporogenous cells is similar in sporangia of the same length from both rows until cell multiplication ceases in sporangia of the megasporangiate row, while it continues in sporangia of the same size in the microsporangiate row. The observed growth differences between sporangia of the micro- and megasporangiate rows are interpreted as events in the differentiation of two sporangial types.     

Apogamous sporophyte formation in a fernPteris multifida and its characteristics

When spores of the fern,Pteris multifida, were aseptically cultured in the dark, sporophytic plants were apogamously induced. The plants have been subsequently grown in pots until the development of leaves with many sporangia for observations of meiotic characteristics in their sporocytes. The sporophytic plants originated from spores were estimated to be haploid, and the estimation was supported by abnormal meiosis in sporocytes and the absence of mature spores, but some chromosomes (n=58) formed bivalents in the meiotic process.     

Phylogeny,Evidence for a Cryptic Plastid,and Distribution of Chytriodinium Parasites (Dinophyceae) Infecting Copepods

Spores of the dinoflagellate Chytriodinium are known to infest copepod eggs causing their lethality. Despite the potential to control the population of such an ecologically important host, knowledge about Chytriodinium parasites is limited: we know little about phylogeny, parasitism, abundance, or geographical distribution. We carried out genome sequence surveys on four manually isolated sporocytes from the same sporangium, which seemed to be attached to a copepod nauplius, to analyze the phylogenetic position of Chytriodinium based on SSU and concatenated SSU/LSU rRNA gene sequences, and also characterize two genes related to the plastidial heme pathway, hemL and hemY. The results suggest the presence of a cryptic plastid in Chytriodinium and a photosynthetic ancestral state of the parasitic Chytriodinium/Dissodinium clade. Finally, by mapping Tara Oceans V9 SSU amplicon data to the recovered SSU rRNA gene sequences from the sporocytes, we show that globally, Chytriodinium parasites are most abundant within the pico/nano‐ and mesoplankton of the surface ocean and almost absent within microplankton, a distribution indicating that they generally exist either as free‐living spores or host‐associated sporangia.     

Induced apogamous sporophytes inPteris dispar andP. semipinnata,and the meiotic behavior in their sporocytes

Spores ofPteris dispar andP. semipinnata were aseptically cultured in flasks for apogamous sporophyte induction. Calli or cell colonies similar to calli were induced in cultures supplemented with hormones. Sporophytic leaves subsequently developed from them in hormone-free medium and the young sporophytes were raised into plants with sporangia. Since the wild-type plants having 116 chromosomes are tetraploid, the sporophytic plants originating from spores would appear to be diploid (dihaploid). In induced sporophytes ofP. semipinnata, non-homologous chromosomes (58 univalents) were found during the meiotic process in sporocytes. InP. dispar, however, the meiotic cells showed many bivalent chromosomes (maximum 29ll). These results suggest thatP. semipinnata is allotetraploid, whereasP. dispar is autotetraploid.     

Re-examination of specimens attributed to Sawdonia curstipa Wang and Hao (zosterophyll) from the Middle Devonian of Xinjiang, China

Specimens previously attributed to Sawdonia curstipa were re-examined. Some of the specimens are transferred to Serrulacaulis cf. S. furcatus based on axial morphology. Other specimens, including fertile axes with one or two rows of sporangia with curved stalks and sterile axes with cone-shaped spines covering the surface, are of uncertain assignment due to poor preservation. The nomenclature problem of Sawdonia custipa is also noted.     

THE SYNANGIATE NATURE OF DOLEROTHECA

Transverse sections through the distal region of a Dolerotheca specimen in a coal ball from the Calhoun coal of Illinois show the double rows of sporangia where they are free from one another. This leads to the conclusion that the sporangial rows should be regarded as extending inward from the campanulum cover rather than radiating outward from the center. This observation suggests that Dolerotheca is a synangium composed of many sporangial units of different lengths.     

CRENATICAULIS,A NEW GENUS OF DEVONIAN PLANTS ALLIED TO ZOSTEROPHYLLUM,AND ITS BEARING ON THE CLASSIFICATION OF EARLY LAND PLANTS

The name Crenaticaulis verruculosus is proposed for slender, pseudomonopodially and dichoto-mously branching plants that bore opposite to subopposite sporangia along the stem and two rows of prominent, multicellular teeth on opposite sides of the terete axis. Epidermal cells were either narrow and elongate parallel to the stem or short with a papillate outer tangential wall. Axillary tubercles were present on one side of the stem near lateral branches. Some tubercles bore remnants of branches. Sporangia dehisced along their distal margins into two unequal halves. No spores were found. Occasional short lengths of stem were petrified by iron pyrite. Sections revealed a cortex consisting of four to six rows of thick-walled cells and a xylem strand. The strand was elliptical in transverse section and maturation was exarch. Tracheids were chiefly scalariform. It is suggested that the so-called axillary tubercles, known in several plants of Devonian age, were scars of rhizophores like those in the modern genus Selaginella. The plant is referred to the subdivision Zosterophyllophytina.     

A new microsporangiate organ from the Lower Carboniferous of the Novgorod Region,Russia

A new species of the genus Telangiopsis, T. nonnae O. Orlova et Zavialova, was described on the basis of a microsporangiate organ from the Lower Carboniferous deposits of the Novgorod Region. The morphology of branching fertile axes, synangia, and sporangia was thoroughly studied. The three-dimensional system of fertile axes branches monopodially; ultimate axes bear numerous connivent bunches of synangia, which consist of three to six basally fused elongated ovate sporangia. The morphology and ultrastructure of prepollen grains were studied, which were extracted from the rock matrix surrounding the sporangia. The two-layered exine includes a well-developed endexine and an alveolate ectexine, with one-three rows of large thin-walled alveolae. The new species was compared with other Early Carboniferous microsporangiate organs.     

The oral apparatus of tadpoles of Rana dalmatina, Bombina variegata, Bufo bufo, and Bufo viridis (Anura)

The morphology and development of the larval oral apparatus of Rana dalmatina, Bombina variegata, Bufo bufo, and Bufo viridis are described and compared using scanning electron microscopy. The species show different arrangements of the mouthparts. The small oral apparatus of R. dalmatina larvae has three labial tooth rows on the upper labium, while there are four tooth rows on the lower labium with a medial gap in row proximal to the mouth. The margins of the oral apparatus are defined by papillae that encircle the lower labium. B. variegata tadpoles have two upper labial tooth rows and three lower labial tooth rows that are uninterrupted, unlike the ones of R. dalmatina. The mouth is encircled by papillae that are larger than those of R. dalmatina. The oral discs of tadpoles of both B. bufo and B. viridis are similar. They are defined by two upper labial tooth rows (the second of which is interrupted by a medial gap) and by three lower tooth rows that differ in lengths in the two Bufo species. Both species develop papillae on the mouth angles and in two rows on the upper labium. Some morphological differences among the oral discs of R. dalmatina, B. variegata, B. bufo, and B. viridis tadpoles can be attributed to phylogenetic differences, but most can be related to their varying feeding habits and/or to their dietary specializations.     

Evolution of apolar sporocytes in marchantialean liverworts: implications from molecular phylogeny

In meiosis of basal land plants, meiotic division planes are typically predicted by quadri-lobing of the cytoplasm and/or quadri-partitioning of plastids prior to nuclear divisions. However, sporocytes of several marchantialean liverworts display no indication of premeiotic establishment of quadripolarity, as is observed in flowering plants. In these cases, the shape of sporocytes remains spherical or elliptical and numerous plastids are distributed randomly in the cytoplasm during meiosis. Through a survey of sporocyte morphology in marchantialean liverworts, we newly report the occurrence of apolar sporocytes in Sauteria japonica and Athalamia nana (Cleveaceae; Marchantiales). Molecular phylogenetic analyses revealed that the quadri-lobing of cytoplasm and quadri-partitioning of plastids were lost independently several times during the evolution of marchantialean liverworts. In addition, our phylogenetic analyses indicate that the simplified sporophytes of several marchantialean liverworts are not a primitive condition but rather represent the result of reductive evolution. The loss of the quadripolarity of sporocytes appears to correlate with the evolutionary trend of the sporophyte towards reductions. Through the evolution of the simplified sporophytes, suppression of mitotic divisions of sporogenous cells might had caused not only the modification of sporophyte ontogeny but also the drastic cytological change of sporocyte.     

DEVELOPMENT OF ENDOPHYTIC FRANKIA SPORANGIA IN FIELD- AND LABORATORY-GROWN NODULES OF COMPTONIA PEREGRINA AND MYRICA GALE

Field-collected nodules of Comptonia peregrina (L.) Coult. and Myrica gale L. (Myricaceae), infected by the nitrogen-fixing actinomycete Frankia sp., were of two types: those that lacked sporangia entirely, designated spore(-), and those that showed extensive sporangial development, designated spore(+). In spore(+) nodules of C. peregrina, sporangia began to develop after the differentiation of endophytic vesicles and the concomitant onset of nitrogenase activity. At the onset of sporangial differentiation, infected host cells appeared healthy. However, endophytic vesicles and host cell cytoplasm and nuclei began to senesce rapidly as sporangia developed. Staining of sectioned material with the fluorescent stain Calcofluor White suggested that vesicles, hyphae and young sporangia were enclosed within a host-derived encapsulation layer, but mature sporangia were no longer encapsulated. In both C. peregrina and M. gale, vesicles were more short-lived in spore(+) than in spore(-) nodules. Field-collected spore(+) M. gale nodules exhibited a pronounced seasonality of sporangial formation. Sporangia began to differentiate in June, after the formation of vesicles and became more prominent in late summer. Inter- and intraspecific cross-inoculations suggest that the ability to form sporangia in the symbiotic state is controlled by endophytic strain type rather than host genotype or host/endophyte combination. The host may, however, influence the number and seasonal appearance of sporangia formed.     

Etude ultrastructurale de l'endoplasme et des vacuoles chez deux types de Dinoflagellés appartenant aux genres Noctiluca (Suriray) et Blastodinium (Chatton)

Résumé L'ultrastructure des éléments cytoplasmiques de deux Dinoflagellés a révélé des différences fondamentales directement liées à leur mode de vie.Chez Noctiluca (Dinoflagellé libre, incolore), les corps de Golgi jouent un rôle important dans la formation des sphères archoplasmiques au moment des divisions et lors de l'élaboration de l'endoglée. Les mitochondries de grande taille, parfois géantes, ont des crêtes nombreuses et une matrice réduite. Elles offrent souvent des relations de contiguïté avec les granulations lipidiques et l'enveloppe nucléaire; elles sont très abondantes dans le tentacule du trophozoïte. Il n'existe pas de plastes. Les inclusions lipidiques abondent ainsi que les trichocystes fibreux et les poches mucifères dont le contenu est polysaccharidique.Chez Blastodinium (Dinoflagellé parasite, coloré sauf une espèce), les corps de Golgi sont très réduits dans le trophocyte, assez rares mais de structure classique dans les sporocytes. Les mitochondries sont de petite taille, pauvres en crêtes et ont une matrice importante chez le trophocyte; elles sont plus grandes dans les sporocytes. Les plastes bien développés, périphériques, possèdent des lamelles stromatiques formées de trois thylakoïdes chez les sporocytes; par contre, chez l'individu végétatif, le plus souvent incolore et relativement protégé de la lumière par plusieurs rangées de sporocytes, les plastes, de grande taille ont un aspect dégénéré: lamelles stromatiques rares, thylakoïdes tassés et stroma important; ce sont des étioplastes. Le cas de Blastodinium contortum hyalinum toujours incolore, est particulier: les plastes existent encore mais sont totalement dégénérés. Il y a très peu de lipides dans le trophocyte, mais ils sont par contre abondants dans les sporocytes. Il existe des trichocystes fibreux et chez l'individu végétatif, des figures myéliniques.Ces importantes variations structurales illustrent parfaitement, dans l'échelle évolutive des Dinoflagellés, l'exemple de deux modes de vie très différents.

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Fine structure of endoplasm and vacuoles in two types of dinoflagellates of the genus Noctiluca and Blastodinium

Summary The fine structure of cytoplasmic elements of two Dinoflagellates reveals fundamental differences directly linked to their way of life.In Noctiluca, free-living, colourless dinoflagellate, the Golgi bodies play an important role in the formation of archoplasmic spheres at the moment of division and when the endoglea is elaborated. The mitochondria of great size, sometimes giant, have very abundant cristae and a reduced matrix. They often show relations of contiguity with the lipid granules and the nuclear envelope; they are very abundant in the tentacle of the trophozoïd. There are no plastids. The lipid inclusions are very abundant and so are fibrous trichocysts and muciferous bags whose content is polysaccharidic.In Blastodinium, parasitic dinoflagellate, coloured except for one species, the Golgi bodies are much reduced in the trophocyte, rather rare but of classical structure in the sporocytes. The mitochondria are of small size, with rare cristae and have an enlarged matrix in the trophocyte; they are bigger in the sporocytes. The plastids are well developed, peripheral, and have stromatic lamellae formed of three thylakoïds in the sporocytes. On the other hand, in the vegetative individual, usually colourless since it is protected from the light by many rows of sporocytes, the large plastids have a degenerated aspect, rare stromatic lamellae, packed thylakoïds and enlarged stroma; they are etioplastids. The case of Blastodinium contortum hyalinum, always colourless, is special: the plastids still exist but are completely degenerated. There are very few lipids in the trophocyte but abundant ones in the sporocytes. Some fibrous trichocysts occur, and in the vegetative individual, some myelinic vacuoles.These important structural variations illustrate very well the example of two very different ways of life, in the evolutionary tree of dinoflagellates.

Cet article fait partie d'un travail d'ensemble constituant une Thèse de Doctorat d'Etat intitulée: La cytologie et la division de deux types de Dinoflagellés: Noctiluca Suriray (genre libre) et Blastodinium Chatton (genre parasite).     

STUDIES OF PALEOZOIC MARATTIALEANS: THE MORPHOLOGY AND PROBABLE AFFINITIES OF TELANGIUM PYGMAEUM GRAHAM

Telangium pygmaeum Graham is known from Upper Pennsylvanian coal balls from the Calhoun coal mine (Illinois). The species was described as possessing radial synangia consisting of 3-5 sporangia fused laterally for about f13 their length. Synangia were believed to be sessile and borne terminally or laterally on a branching rachis without lamina. Examination of new coal ball material of the same age indicates that the synangia are borne abaxially on the pinnules of a compound frond with the anatomy of a Psaronius leaf (Marattiales). Synangia are sessile and borne in two rows, one on either side of the pinnule midrib, under the unbranched lateral veins. Synangia are radial, 0.6 mm in diam, and consist of a ring of thin-walled sporangia fused to near their apices prior to dehiscence, but separating on dehiscence to release spores along their inner midline. Spores are spherical, trilete, 30-48 μm in diam, with a granulate ornamentation. The new genus Araiangium is proposed for this material based on the organization of the sessile thin-walled synangia. Araiangium is compared with other marattialean genera with sessile synangia (Acaulangium, Acitheca), and with the pedicellate synangia of various species of Scolecopteris. Criteria used in the delimitation of genera in Paleozoic anatomically preserved marattialean fertile foliage are discussed.     

Parthenogenesis is rare in the reproduction of a sexual field population of the isogamous brown alga Scytosiphon (Scytosiphonaceae,Ectocarpales)

Parthenogenetic development of unfused gametes is commonly observed in laboratory cultures among various brown algal taxa. There is, however, little information on the contribution of parthenogenesis to the reproduction of field populations. In this study, we investigated whether parthenogenesis is present in a sexual population of the isogamous brown alga Scytosiphon with a 1:1 sex ratio. In culture, both female and male gametes showed higher mortality and slower development compared to zygotes. More than 90% of surviving partheno‐germlings formed parthenosporophytes irrespective of the culture conditions tested. Therefore, if parthenogenesis occurs in the field, most unfused gametes are expected to form parthenosporophytes. Contrary to this expectation, parthenosporophytes were rare in the field population. We collected 126 sporophytic thalli and isolated and cultured a unilocular sporangium from each of them. We confirmed that cultures of 120 unilocular sporangia produced both female and male gametophytes by the observation of zygotes or amplification of PCR‐based sex markers indicating that these sporangia originated from zygotic sporophytes. Only females were detected in cultures from two sporangia and only males from four sporangia suggesting that these sporangia originated from parthenosporophytes. In the Scytosiphon population, although parthenogenesis is observable in culture, our results demonstrate that the contribution of parthenogenesis to reproduction is small (≤4.8%) compared to sexual reproduction. Unfused gametes may not survive to form mature parthenosporophytes in significant numbers in the field partly due to their higher mortality and slower development compared from zygotes.     

A spectrophotometric approach for determining sporangium and zoospore viability of Plasmopara viticola

A spectrophotometric method for determining the viability of sporangia and zoospores of the oomycete Plasmopara viticola (causal agent of grapevine downy mildew) is described and evaluated to overcome the limitations of currently available methods for assessing propagule viability. Sporangia produced on leaf discs in the laboratory were harvested at different days after the initiation of sporulation (DAS) to yield differences in sporangium viability. Sporangia were suspended in sterile water, the suspensions were placed in a cuvette, and sporangium germination was monitored in a spectrophotometer (λ = 600 nm) at 2- to 3-min intervals for 5 hr. Absorbance started to increase after sporangia were suspended in water for ~30–60 min followed by major peak(s) for younger sporangia (1–3 DAS), whereas low to no increase in absorbance was observed for senescent sporangia (>7 DAS). Microscopic observation confirmed that the increase in absorbance corresponded to the release and active swimming of zoospores, whereas absorbance decreased when zoospores encysted and settled. A positive correlation (r = .839, p = .0365) was observed when the time to the initial increase in absorbance was plotted against the age of sporangia. The time to the absorbance peak (marking the time of maximum zoospore movement) was shortest for immature sporangia (0 DAS), longest for young sporangia (2 DAS) and decreased for mature and senescent sporangia. A similar pattern was observed for the standardized area under the absorbance curve (indicating the overall quantity of zoospores released), for which values were lowest for immature and senescent sporangia, highest for young sporangia and intermediate for mature sporangia. Consistent patterns obtained across two independent experiments suggest that the method is reproducible and may be further developed for other zoospore-releasing pathogens.     

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。     

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.     

Ontogeny of strobili, sporangia development and sporogenesis in Equisetum giganteum (Equisetaceae) from the Colombian Andes

Studies on the ontogeny of the strobilus, sporangium and reproductive biology of this group of ferns are scarce. Here we describe the ontogeny of the strobilus and sporangia, and the process of sporogenesis using specimens of E. giganteum from Colombia collected along the Rio Frio, Distrito de Sevilla, Piedecuesta, Santander, at 2200m altitude. The strobili in different stages of development were fixed, dehydrated, embedded in paraffin, sectioned using a rotatory microtome and stained with the safranin O and fast green technique. Observations were made using differential interference contrast microscopy (DIC) or Nomarski microscopy, an optical microscopy illumination technique that enhances the contrast in unstained, transparent. Strobili arise and begin to develop in the apical meristems of the main axis and lateral branches, with no significant differences in the ontogeny of strobili of one or other axis. Successive processes of cell division and differentiation lead to the growth of the strobilus and the formation of sporangiophores. These are formed by the scutellum, the manubrium or pedicel-like, basal part of the sporangiophore, and initial cells of sporangium, which differentiate to form the sporangium wall, the sporocytes and the tapetum. There is not formation of a characteristic arquesporium, as sporocytes quickly undergo meiosis originating tetrads of spores. The tapetum retains its histological integrity, but subsequently the cell walls break down and form a plasmodium that invades the sporangial cavity, partially surrounding the tetrads, and then the spores. Towards the end of the sporogenesis the tapetum disintegrates leaving spores with elaters free within the sporangial cavity. Two layers finally form the sporangium wall: the sporangium wall itself, with thickened, lignified cell walls and an underlying pyknotic layer. The mature spores are chlorofilous, morphologically similar and have exospore, a thin perispore and two elaters. This study of the ontogeny of the spore-producing structures and spores is the first contribution of this type for a tropical species of the genus. Fluorescence microscopy indicates that elaters and the wall of the sporangium are autofluorescent, while other structures induced fluorescence emitted by the fluorescent dye safranin O. The results were also discussed in relation to what is known so far for other species of Equisetum, suggesting that ontogenetic processes and structure of characters sporoderm are relatively constant in Equisetum, which implies important diagnostic value in the taxonomy of the group.     

BARINOPHYTON CITRULLIFORME (BARINOPHYTALES INCERTAE SEDIS,BARINOPHYTACEAE) FROM THE UPPER DEVONIAN OF PENNSYLVANIA

Barinophyton citrulliforme is definitely proven to be a tracheophyte. The vascular cylinder of the main axis is an exarch protostele composed of tracheids having a continuous secondary wall folded into protrusions into the cell lumen. These protrusions delineate the position of annular thickenings which were deposited earlier than the continuous secondary wall. Between successive protrusions, the later-deposited secondary wall is interrupted by minute pitlike structures. It is suggested that the secondary walls of the tracheids were laid down in a two-phase depositional sequence. The fertile system of B. citrulliforme consists of a main axis bearing spirally arranged strobili. The strobilus consists of an axis that bears two alternate rows of sporangiferous appendages. The sporangiferous appendages are borne laterally along the strobilar axis and recurve abaxially around the axis. The sporangia are attached along the inside curve of the appendages, one sporangium per appendage, each containing both microspores and megaspores. This species thus exhibits sporangial heterospory which is considered to be an adaptation to an aquatic habit and the sporangia are considered to be functional analogs of the sporocarps of Marsilea. The interpretation of the strobilus is morphologically identical to Ananiev's interpretation of the strobilus of Pectinophyton bipectinatum; consequently P. bipectinatum is here transferred to Barinophyton as B. robustius comb. nov.