VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS. IV. OUTER SURFACE MUCILAGE OF NAVICULA CONFERVACEA VAR. CONFERVACEA1

The development of the mucilage on the outer surface of Navicula confervacea (Kütz.) Grun., a raphed, filamentous diatom, was studied with scanning electron microscopy. This nonstructural cell wall material, present on the surface after critical-point drying and absent after acid cleaning, was of two types: strands and papillae. Strands were associated with the raphe system, areolae, elongated pores of the mantle, and all girdle sutures. Organic papillae were a common feature of valves, valvo-copulae and pleurae, but their origin and distribution could not be explained since they often occurred between the obvious openings in the frustule. Strands from the raphe and areolae may function in attaching terminal cells to a substrate and adjacent cells to each other. Other strands of the girdle arise from sutures during cell enlargement and continue to lengthen and intertwine until the individual frustules within a filament are obscured. Strands from sutures might originate from the advalvar row of pores of the girdle bands since these pores lie along the suture, but direct observation of this was not made. Secretion between, the bands also cannot be ruled out. Although mucilaginous papillae may sometimes occur at random on the entire surface of frustules, there is also a distinct, narrow multiseriate row of them around the edge of valves without marginal spines.     

Eolimna martinii n. gen., n. sp. (Bacillariophyceae) aus dem Unter-Oligozän von Sieblos/Rhön im Vergleich mit ähnlichen rezenten Taxa

The laminated sediments of the Lower Oligocene ‘Sieblos-Schichten’ (dysodile and kieselgur-sediments) contain many individuals of a naviculoid diatom species. Formerly this group of diatoms would have been classified without doubt as Navicula (section ‘Minusculae’). After splitting this extremely heterogeneous genus in Navicula sensu stricto and some other homogeneous genera, the pattern of structures of this fossil species does not correspond to any established genus. For that reason this form will be described as the generotype of a new genus Eolimna. Furthermore it became evident that some Recent species (assigned to Navicula) show the same pattern of structures as the new genus Eolimna (generotype: E. martinii) which represents one of the first naviculoid diatoms of limnic origin. The essential criteria of this genus are: comparatively small cells with narrow girdle; simple alveolated rib system; areolae with a hymen in approximately medium position between relatively large sized foramina outside and inside of the valve; a single row of more or less irregular arranged areolae in close position to each other, on one or several of the copulae. The differences to supposed related genera are discussed.     

STRUCTURE,LIFE HISTORY AND SYSTEMATICS OF RHOICOSPHENIA (BACILLARIOPHYTA). I. THE VEGETATIVE CELL OF RH. CURVATA1

Rhoicosphenia Grun. has been placed by some authors in the monoraphid group with Achnanthes Bory and Cocconeis Ehrenb., and by others near Gomphonema Ehrenb. In order to clarify the systematic position of the genus, the morphology and anatomy of the vegetative cells of Rh. curvata (Kütz.) Grun. were investigated using light and electron microscopy. The structure and formation of the two types of valve are described, and the heterovalvy shown to be of a different type from that of the monoraphids; on the basis of raphe, valve and girdle structure a close relationship between these and Rhoicosphenia is unlikely. Rhoicosphenia shows many resemblances to Gomphonema but the types of pore occlusion present, coupled with apparently slight differences in the mucilage-secreting structures and the girdle, suggest that classification in the same family is unwise. The cryptic asymmetry of the valves, and in particular of the raphe system, is noted and explained with reference to their formation; with respect to this asymmetry two configurations of the valves can occur (named cis and trans types) and the distribution of these in raphid genera is discussed briefly. In view of the lack of evidence in raphid diatoms supporting a classification of bands into copulae and pleurae, it is recommended that this practice be suspended.     

THE MORPHOLOGY AND INTEGRATION OF VALVES AND BANDS IN NAVICULA MUTICA VAR. MUTICA (BACILLARIOPHYCEAE)1

Navicula mutica (Kütz.) var. mutica was isolated from the air, cloned on agar, cultured in soil-water bottle, and studied with transmission and scanning electron micros-ropy. The frustules were lanceolate to ovoid with rounded apices, with the apical axis 8.5 ± 3.2 μ and the trans-apical and the transapical axis 3.6 ± 0.6 μm. Striae were composed of two or three puncta, and the mantle bore a single row of puncta aligned with the striae. The ends of the raphe turned away from an isolated punctual in the central area of the valve. The mantle puncta and one or two of the valve-face puncta in each stria opened into a series of transapical grooves in the interior of the valve, the grooves contributing to the appearance of striae in the light microscope. The interior of the mantle also possessed a pair of longitudinal grooves, discontinuous at the apices of the valves. An undulate advalvar margin of the valvocopula likely articulates along the interior longitudinal groove of the mantle. The projections of the undulate margin are perhaps positioned between the transapical grooves and along the longitudinal groove between the dentiform structures formed by the intersection of the double-grooved system. The girdle bands each had two (occasionally three) rows of pores. The pleurae margins were straight and not undulate.     

VARIATION IN PATTERNS OF VALVE MORPHOGENESIS BETWEEN REPRESENTATIVES OF SIX BIRAPHID DIATOM GENERA (BACILLARIOPHYCEAE)

Scanning electron microscopic studies of silica valve formation in naviculoid diatoms representing six different genera revealed that the precise sequence of depositional events varied among genera. Valve deposition begins with the formation of the raphe sternum, from which virgae (lateral outgrowths) extend. Areolae (pores) are formed between the virgae by the fusion of cross-extensions (vimines). In most of the species studied ( Craticula ambigua (Kützing) D. G. Mann, Frustulia vulgaris (Thwaites) De Toni, Craspedostauros australis E. J. Cox, and Gomphonema truncatum Ehrenberg), areola (pore) formation began near the raphe sternum before completion of the valve margin, but in Pinnularia gibba Ehrenberg the valve margin fused before the areolae were formed. Silica deposition in all these taxa was mainly distal to proximal (with respect to the cytoplasm), but in Haslea sp. it was mainly proximal to distal. Haslea also differed in that areolae were defined as the valve margin was completed. These data have also contributed to the interpretation of taxonomically important features, such as raphe endings. In P. gibba the internal central raphe fissures were laterally deflected but subsequently obscured by additional silicification of the valve, whereas in G. truncatum they were initially straight, becoming laterally deflected as valves mature. External raphe fissures in Frustulia became Y-shaped only just before maturity; in immature valves they were dotlike, as in Amphipleura Kützing. The comparison of developmental pathways in diatoms is a useful adjunct to morphological and other approaches in diatom systematics and warrants renewed attention.     

Morphology of the marine epiphytic diatom Cocconeis heteroidea (Bacillariophyceae)

The morphology and fine valve structure of the marine epiphytic diatom Cocconeis heteroidea Hantzsch have been investigated. The entire frustule, including the internal and external structure of the raphid valve (RV) and araphid valve (AV), and the complete cingulum, are described using light microscopy and scanning and transmission electron microscopy, using a bleaching method. The strongly sigmoid raphe terminates in elongate hooked helictoglossae internally. The hymenes, with perforations arranged in a centric array, are located near the internal openings of the areolae in the RV. The striae in the AV consist of alveoli occluded by hymenes, that have perforations arranged in a parallel array and are located near the outer surface. The complete cingulum of AV consists of three open bands without fimbriae: a valvocopula, a copula with a ligula and a pleura with a small ligula. The RV has only a valvocopula which is open type and not fimbriate.     

The transfer of Fragilaria obtusa Hustedt to the genus Staurosira Ehrenberg (Bacillariophyceae)

The ultra‐structure of the frustule of the rarely recorded diatom Fragilaria obtusa Hustedt was studied in detail using sand samples from Laranjal Bay (Lagoa dos Patos Rio Grande do Sul State, Brazil). F. obtusa is transferred to the genus Staurosira Ehrenberg as Staurosira obtusa (Hustedt) Garcia. The taxon is characterized by striae composed of elliptical areolae that are occluded by an internal velum, apical pore field formed by several rows of rimmed pores, girdle bands free of ornamentation, wide valvocopulae, and absence of rimoportulae. This is the first record of S. obtusa from an epipsammic habitat.     

Cover

ON THE COVER: Valve formation in a raphid diatom Diploneis smithii observed under SEM. The initial 2D network of ribs (top) thickens and begins vertical development (middle) to form an elaborate 3D structure with alternately arranged areolae, elliptical margin, and longitudinal canals on either side of the midrib containing the raphe slits. All valves are shown from the inside. [Vol. 54, No. 2, pp. 171–186 ]     

VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS. II. INTEGRATION OF VALVES AND BANDS IN NAVICULA CONFERVACEA VAR. CONFERVACEA12

Chemically cleaned and critical-point dried cells of a clonal culture were examined with scanning electron microscopy. Cells form filaments by valve-to-valve connections maintained by organic material which adheres to the central area of the valve face. Bending of filaments is probably restricted to some extent by the articulation of overlapping spatulate marginal spines with an adjacent underlapping set of much shorter spines (ridges), and with the mantle edge itself. Cell division results in three possible spine patterns for each cell: a set of overlapping and a set of underlapping spines; no overlapping sets of spines (two underlapping); or two sets of overlapping spines (no underlapping). Each filament inherits cells with spine set patterns in the ratio of 2 (with 1 set overlapping): 1 (with no sets overlapping): 1 (with 2 sets overlapping). Valvocopulae are shaped similarly to pleurae except that the partes exteriores of the valvocopulae are wider. The pars interior of both is delimited by an advalvar row of pores continuous around the cell apex. The pars exterior also has a row of pores, but it is median in the valvocopula and first pleura and does not continue around the cell apex. The valvocopulae always underlap the mantle and the pleurae always underlap their preceding band. The ends of both appeared attached, but may become free in acid-cleaned preparations. Bands alternate with each other so that the ends of the valvocopula attach to the first continuous apical portion of the first pleura; the ends of the first pleura attach in that same fashion to the second pleura but at the opposite apex; and all subsequent pleurae alternate in the same fashion with up to at least 13 pleurae/epicingulum. The continuous apical portion of each band is elevated so that a functional (but not structural) ligula is formed, with the continuous apical portion of alternate bands becoming adjacent and underlapping each other only in this region. The valvocopulae in a single cell, or of adjacent cells, may have their continuous apical ends on the same or on opposite apices. It is recommended that N. confervacea var. peregrina (W. Sm.) Grun. be merged with the nominate variety.     

Studies on the biochemistry and fine structure of silicia shell formation in diatoms VII. Sequential cell wall development in the pennateNavicula pelliculosa

Summary The development of the wall of synchronized culture ofN. pelliculosa is described. The first step, modification of the 3-2 configuration of the girdle bands of the wall during interphase, occurs immediately before mitotic division by the addition of a third girdl band to the hypotheca. Following cytokenesis, the new valve is initiated when a primary central band is formed within a silica deposition vesicle. This band extends the length of the cell and contains a central nodule. Secondary arms extend from the central nodule, join with extensions of the primary central band, and constitute the raphe rib. Mounds or knolls are formed on the central nodule and disappear as the valve matures. Transapical ribs appear on both the primary central band and secondary arms, and cross extensions join to form the sieve plate areas. The wall appears to be released by a joining of the inner silicalemma and the plasmalemma. An organic coat covers the newly released wall. Two girdle bands are formed and released sequentially.     

WALL MORPHOGENESIS IN COSCINODISCUS WAILESII GRAN AND ANGST. I. VALVE MORPHOLOGY AND DEVELOPMENT OF ITS ARCHITECTURE1

The morphology of the valve of Coscinodiscus wailesii and the development of its siliceous architecture, studied in the SEM and TEM, is compared with valve formation in Thalassiosira eccentrica (Ehrenberg.) Cleve. Though the areolae-architecture of these two species differs in such that the cribrum is proximal and the foramen distal in T. eccentrica, and in C. wailesii the cribrum is distal and the foramen proximal, the formation of their complex loculate system is similar, displacing a centrifugal growth pattern with respect to the valve, and a proximal to distal, sequentially. During base layer formation a hitherto undescribed rib system outlines the prospective areolae. The vertical differentiation is in principle the same as in T. eccentrica and also the cribra are formed centripetally in relation to the areolae in both species. The location of the cribra at the proximal or distal side, therefore, seems to be of minor importance for the sequence of silica deposition. Variation in girdle bands is discussed with respect to cell division. The prophasic nuclear migration from the interphase position to the girdle bands, where mitosis is performed, seems to be necessary for triggering the formation of the unilateral cleavage furrow that later forms a cleavage ring with excentric position. The divided nuclei migrate with the ingrowing cleavage furrow to the center of the newly created protoplasmic surfaces to initiate valve formation.     

SEA ICE DIATOMS (BACILLARIOPHYCEAE) OF THE CANADIAN ARCTIC. I. THE GENUS STENONEIS1

Ice diatom assemblages in the Canadian Arctic are dominated by the Naviculaceae, a family which includes the genus Stenoneis Cleve. A taxonomic history of S. inconspicua var. baculus (Cleve) Cleve (including type material of Navicula baculus Cleve) is presented with morphological data including the first SEM views of the genus. Stenoneis inconspicua var. baculus was compared with S. obtuserostrata (Hustedt) comb. nov. (basionym: Navicula obtuserostrata Hustedt). In the SEM, external, short, flattened siliceous ridges were revealed beside the proximal parts of the raphe branches and central pores. This may be a significant taxonomic feature of the genus. Valves of S. inconspicua var. baculus ranged from 40–103 μm in length, 7–11 μm in width and had 17–20 striae in 10 μm, whereas valves of S. obtuserostrata were shorter (27–53 μm), narrower (6–7.5 μm) and more finely striated (24–30 in 10 μm). Both taxa have a circumpolar Arctic distribution.     

Valve morphogenesis in an araphid diatom Rhaphoneis amphiceros (Rhaphoneidaceae,Bacillariophyta)

The pattern centre in valve morphogenesis is an annulus in centric diatoms or a sternum in pennate diatoms. The genus Rhaphoneis is currently placed within a lineage that diverges at the root of the pennate diatom clade in most molecular phylogenies, and its valves have a unique pattern to their striae, i.e. radiating from both apices, giving the impression that a pattern centre exists at both ends of the valve and virgae (ribs) formation proceeds centripetally. The present study, however, shows that the pattern centre is actually a linear sternum and the formation of virgae proceeds centrifugally, a pattern centre that is commonly found in most araphid diatoms. Thus, the hypothesis that valve morphogenesis based on a linear sternum and perpendicular virgae is a synapomorphy of pennate diatoms is supported. Our study also demonstrates that the pattern of valve formation can be observed by light microscopy with a direct mounting method when the specimen is relatively large, i.e. exceeding approximately 50 µm in valve length. An important advantage of the use of the direct mounting method is that it requires no repeated centrifugation steps for dehydration, steps necessary for observation by a scanning electron microscope, causing the loss and/or collapse of the specimen, particularly with fragile valves in the early stages of development.     

THALASSIOSIRA ANDAMANICA SP. NOV. (BACILLARIOPHYCEAE), A NEW DIATOM FROM THE ANDAMAN SEA (THAILAND)

A new marine diatom, Thalassiosira andamanica, is described from light and electron microscopy. The specimens were collected in the vicinity of Phuket Marine Biological Center, Thailand, and later brought into clonal culture. Thalassiosira andamanica possess a rimoportula with a pronounced outer extension, one marginal ring of fultoportulae, and three rings of fultoportulae on the valve face. Cells are united into colonies by a single thread secreted through a central fultoportula. Marginal fultoportulae extensions are shortest on the inside of the valve. The areolae are arranged in sectors, and the valve margin is ribbed with approximately 38 ribs in 10 μm. The valvocopula and copula have rows of pores, four to six pores in 1 μm. Apparently, the pleurae are hyaline. Experiments with a clonal culture isolated at Phuket, Thailand, showed that growth (cell divisions·24 h⁻¹) was reduced for cultures grown at 14° C compared to those grown at 19°, 24°, or 30° C. The maximum growth rate (2.2 divisions·24 h⁻¹) was at 30° C. Thalassiosira andamanica is compared with morphologically similar taxa. On the basis of morphological features and the response to different temperature regimens, it is concluded that this taxon must be recognized as a new warm-water species. In addition, T. andamanica does not clearly belong to any of the two subgroups of species of Thalassiosira. To accommodate the morphological characteristics of T. andamanica, the establishment of a possible third subgroup is discussed.     

管壳缝类硅藻中国新记录属种——非洲南氏藻

利用光学和电子显微镜对采自黄海水域的1个管壳缝类硅藻——非洲南氏藻进行了形态学研究,并对其地理分布进行了讨论.结果表明:(1)该种壳体带面呈矩形,壳面窄椭圆形,具有钝圆的末端.(2)壳缝居中,由两条等长的分支组成.(3)管壳缝由复杂、接合的肋突支撑,但无龙骨.(4)每条线纹仅有1个孔纹,壳套上最多有1列孔纹.(5)目前...     

THE “PARADOX” DIATOM BACILLARIA PAXILLIFER (BACILLARIOPHYTA) REVISITED1

Defined by its unique colonial locomotion, Bacillaria paxillifer (O. F. Müll.) Hendey was recognized as a single, pandemic species by many phycologists. However, reinvestigation of colonies from different habitats revealed three distinct groups: (A) brackish/freshwater, (B) marine littoral, and (C) marine planktonic taxa. Groups differed in colony and cell form, raphe flanges (RFs), shape and position of transapical ribs (Tr's), and morphogenesis. Linear‐shaped species were restricted to group A: Tr's thickened principally to the interior. Lanceolate forms were confined to groups B and C: valve formation proceeded from an internal base layer to the exterior. The planktonic species differed in the shape of its raphe slit, and the transformation of girdle bands (GBs) into “winglets.” Taxa also differed in chloroplast shape and number. All species formed motile colonies. Siblings adhered via elastic fibrils secreted through their raphe. Raphe ribs were held in position by siliceous clamps (fibulae), anchored in an extra pair of axial ribs (fibular ribs) parallel to the raphe ribs. This raphe system resembled that of Cylindrotheca rather than the “canal raphe” of Nitzschia. Many valves were asymmetric along the apical axis due to protruding RFs shuttling in a 1:1 ratio within a colony, but raphe slits were mirror images, as were the growth direction of fibulae and position of plastids, with pyrenoids tilted in the same direction. Species possessed four open GBs per epitheca; the third band invariably bore an internal, organic ridge to aid in adhesion of the plasmalemma during cleavage. The results suggested that these taxa are a natural phylogenetic group, requiring precise determination of their taxonomic position.     

THE MARINE,PLANKTONIC DIATOMS THALASSIOSIRA OCEANICA SP. NOV. AND T. PARTHENEIA1

The diatom clone 13–1 isolated from the Sargasso Sea by Dr. R. R. L. Guillard as Cyclotella nana Hust. or Thalassiosira pseudonana Hasle and Heimdal is described as a separate species, Thalassiosira oceanica sp. nov. An amplified diagnosis of Thalassiosira partheneia Schrader is given, and a comparison of the two species is made based on light and electron microscopy. Similarities are present in the apparent distribution pattern, cell size, and girdle structure. Differences are present in the shape of the areolae of the valvocopula and the copula, in the structure of the vela of these bands, in the texture of the external valve surface, in the morphology of the strutted processes, in the location of the labiate process, in the distance between the marginal strutted processes, and by the presence of a marginal ridge in T. oceanica. Fifteen nanoplanktonic (maximum diameter = 20 UmUm) Thalassiosira species are listed, among them T. oceanica and T. partheneia.     

SURFACE MORPHOLOGY OF THE MARINE DINOFLAGELLATE SINOPHYSIS MICROCEPHALUS (DINOPHYCEAE) FROM A MANGROVE ISLAND. TWIN CAYS. BELIZE1

Sinophysis microcephalus Nie and Wang 1944 is a nonphotosynthetic, tropical, benthic. dinophysoid dinoflagellate. I isolated it from floating detritus on a subtropical mangrove island. Twin Cays. Beleze, Central America, and describe its micromorphology from light and scanning electron micrographs. Cells of S. microcephalus are circular to subcircular and compressed laterally with a cell size of 42-44 μm long and 33–35 μm wide and with a length /width ratio of 1.25–1.28. Areolae are numerous, 368–550 per valve, ranging in size from 0.75 to 2.0 μm. Pores are oblong and deeper at the valve's center and pentagonal-shaped at the plate margin. The well-defined cingulum is narrow and deeply incised with a smooth surface. The epitheca is small, moderately convex, and divided into two large, highly ornate, asymmetrical plate: the left and right epitheca I plates. The left epithecal plate bears two slightly curved, upright anterior projections located dorsally adjacent to the epithecal list, a relatively large opening, and three smaller openings compressed against the sagittal suture. The right plate contains a wide megacytic zone with two parallel ridges, a fairly large oblong opical pore in ventral position adjacent to the cingulum, and eight areolae each with a round, uniform-sized pore opening. There are two long and narrow sulcal lists, gently convex with a smooth edge without structure or ribs. The left sulcal list has an ear-shaped labe, a form of a primitive dinophysoid list. The megacytic zone is smooth and expands unevenly during cell division. The epitheca and sulcus distinguishes S. microcephalus from all examined Dinophysis.     

VALVE MORPHOGENESIS AND THE MICROTUBULE CENTER IN THREE SPECIES OF THE DIATOM NITZSCHIA1

The relationship of cell organelles to valve morphogenesis was investigated in three species of Nitzschia. One, N. sigmoidea (Nitzsch) W. Sm., showed consistent ability to generate both nitzschioid and hantzschioid symmetry in daughter cells following cytokinesis; the other two maintained nitzschioid symmetry stably. From previous work with Hantzschia, a certain sequence of events could be anticipated in the cytoplasm. In two significant areas–the behavior of the Microtubule Center (MC) and its microtubule (MT) system, and the central origin of the silicalemma–not only were the results unexpected, but the three species showed fundamental differences among themselves. In N. sigmoidea, the silicalemma (and the future raphe region) arises centrally on the cleavage furrow, and after some lateral expansion, the silicalemmas and their associated organelles move in opposite directions in daughter cells, so that the raphe and the raphe canals end up along the girdle side of the cell as expected. However, the MCs never become associated with their silicalemma, remaining throughout near the girdle bands. In N. sigma (Kütz) W. Sm., the silicalemmas arise centrally and after lateral growth, move in opposite directions to generate nitzschioid symmetry. In this case, the MCs move to the vicinity of but never close to the silicalemmas, and follow them distantly during their lateral movement. In N. tryblionella Hantzsch, the new silicalemmas arise opposite one another, on one side of the daughter cells; each MC soon moves very close to its silicalemma, and remains thus through most of valve morphogenesis. Later, only one silicalemma/MC complex moves laterally, establishing the nitzschioid symmetry in both daughter cells. In all three species, as in Hantzschia, linear arrays of mitochondria aligned along MTs occupy the forming raphe canal, and microfilaments line the outer edge of the expanding silicalemma. The fibulae (the wall struts arching across the raphe canal) in Hantzschia always grow from the valve surface to the girdle surface of the forming valves. In these three Nitzschiae, this invariably happens in only one daughter cell of any pair; in the other, all the fibulae grow from the girdle surface to the valve surface. An explanation of these variations is proposed: that the morphogenetic machinery of Nitzschia and Hantzschia have a common origin, with present Nitzschiae having undergone considerable diversification at the intracellular level, causing the unstable cell symmetry exhibited by several modern species. Perhaps a taxonomic distinction between Hantzschia and Nitzschia lies in whether the morphogenetic machinery associated with valve morphogenesis moves laterally in the same or in opposite directions.