Organic chemistry of Protosalvinia (Foerstia) from the Chattanooga and New Albany Shales

Vertical samplings of Protosalvinia, a thalloid Upper Devonian alga from the Chattanooga and New Albany Shales, are chemically analyzed and correlated with the organic chemical constituents isolated from associated shale matrices. Normal, saturated acids (n-C₈ to n-C₃₆n-paraffins (C₁₀ to C₃₆), showing an odd carbon-number preference, branched-chain alkanes, and vanadyl porphyrins isolated from Protosalvinia vary in their concentrations with depth of burial and with the dominant associated morphology of Protosalvinia, i.e., P. arnoldii, P. ravenna, P. furcata. Organic constituents of shales, in general, reflect those detected in thalli; relative concentrations, molecular diversity, carbon chain-lengths and maxima of compounds extracted from both shale and fossil material are similar. Pristane, phytane and porphyrins are probably derived from a chlorophyllous organism, while δ¹³C data corroborate a photosynthetic system operating in the primary biosynthesis of shale geochemistry. Crude-oil extracts of Protosalvinia-rich strata contain higher alkane and lower aromatic hydrocarbon concentrations than those of an average crude oil. Chemical variations among forms of Protosalvinia suggest biochemical differences in original plant composition rather than diagenetic transitions; field observations of morphological trends seen in vertical samplings may be used in crude extrapolations of the organic chemical compositions of shale strata.     

STIGMARIAN PETRIFACTIONS FROM THE PENNSYLVANIAN OF COLORADO

The designation Stigmaria ficoides is restricted to casts and impressions. Petrifactions of Stigmaria arnoldii sp. n. occur in Pennsylvanian age strata in Trout Creek Pass, Colorado. The rootlet gaps of this species appear narrow and elongate in both cross and longitudinal sections. Rootlet traces are contiguous with the secondary xylem to its outer margin and are surrounded by a small parenchyma sheath in the middle cortex. The outer cortex consists of an inner layer of primary tissue and a relatively thick zone of periderm. S. arnoldii most closely resembles material described as S. ficoides from England. Various stigmarian characters are evaluated with regard to their systematic or ontogenetic significance.     

A new species of Pleuridium from Australia

Abstract

A new species, Pleuridium curvisetum, is described. It has an arched seta similar to that of P. arnoldii and was collected, sometimes in association with that species, in alpine country in Australia. P. arnoldii was first recorded from New Zealand. A comparison of the two species revealed some features, including a bistratose margin to the leaf subula, not previously reported for P. arnoldii.     

OBSERVATIONS ON THE FINE STRUCTURE OF CALLIXYLON WOOD

Secondary wood of three species of Callixylon of Lower Mississippian age, preserved by three different modes (fusinization, silicification, and phosphatization), have been studied and characterized in detail. Problems of interpretation at the SEM level of permineralized woods, both containing (silicified wood of Callixylon erianum) and essentially lacking (phosphatized wood of C. arnoldii) original organic cell wall material, are analyzed and discussed. In particular, it is concluded that the flat to curved surfaces showing no evidence of apertures, observed in bordered pit pairs, commonly represent pit membrane surfaces. It is accepted, however, that some concave surfaces might be the mineral accretion surfaces of incomplete pit cavity casts as proposed by Leo and Barghoorn (1976). Regions between groups of pits, previously interpreted as crassulae, may be artifacts of preservation. The fusinized wood has the general appearance of charcoal, but unlike commercially produced charcoal provides evidence of its original microfibrillar structure. The origin of fusain in the fossil record is discussed, and it is concluded that it probably had several origins, including forest fire. Since charcoal can be produced naturally in the absence of O₂ (Brown and Davis, 1973), the suggestion that fusain (charcoal) in the geologic column provides a basis for “assessing oxygen levels in paleoatmospheres” (Cope and Chaloner, 1980) is not supported. Natural sites of fusain production in the absence of O₂ are regions of vulcanism and organic sediments inhabited by anaerobic microorganisms. A circular pattern of crystal orientation in the pit borders of C. arnoldii is interpreted to represent the original microfibrillar pattern. Pit apertures in C. arnoldii are shown to be circular to slightly elliptical. Interpretive evidence of heterogeneous pit membranes in C. arnoldii suggests but does not prove the presence of a torus. The distinctive central region in some pit membranes of the fusinized wood of Callixylon sp. might represent accumulations of waste metabolites. It is argued that a torus would be highly adaptive in large pits with circular apertures.     

MORPHOLOGY AND TAXONOMY OF HIMANTOTHALLUS (INCLUDING PHAEOGLOSSUM AND PHYLLOGIGAS), AN ANTARCTIC MEMBER OF THE DESMARESTIALES (PHAEOPHYCEAE)1

The sporophyte of Himantothallus develops according to a closed pattern in which the number and position of the blades is determined by the location of trichothallic meristems in a filamentous germling. Expansion of the miniature juvenile to the massive adult thallus is accomplished by diffuse secondary growth and involves a change from filamentous rhizoids to a hapteroid holdfast, flattening of the stipe, and enormous increases in length, breadth, and thickness of both stipe and blade. The axis usually bears 1–8 lateral blades, often paired, and terminates in a flattened stub. Phaeoglossum is interpreted as a growth form of Himantothallus in which a terminal blade develops to the exclusion of lateral blades, the latter being represented by a single spine. Phyllogigas clearly falls within the morphological spectrum of Himantothallus, the lack of twisting being related to physical factors in the environment. Sporangia, interspersed with an equal or somewhat larger number of two-celled paraphyses, are borne in slightly elevated sori scattered over both surfaces of the blade. Zoospore germination was not observed, nor were gametophytes, either in culture or in the field. Haptera apparently originate from the meristoderm in the lower part of the maturing stipe and lack a filamentous medulla. The mature stipe and the mature blade are anatomically similar, being composed of a superficial meristoderm, a cortex of parenchyma-like cells, and a filamentous medulla. The meristoderm is usually a single layer of plastid-containing cells that divide anticlinally to accommodate (or effect) expansion and periclinally to produce cortical tissue inward. Cortical cells are in radial files and increase in diameter towards the interior. They usually are densely packed with physodes. The medulla is uniquely distinguished by the presence of sheathed trumpet hyphae. Cells of the trumpet hyphae have perforate end walls with callose deposits and probably function in conduction as do the sieve filaments in Laminariales. Sheathing cells are filled with plastids. Sheathing filaments form connections among themselves and with nearby unsheathed filaments. The sheathed trumpet hyphae and their matrix of unsheathed filaments form a plexus, which in the mature blade is flattened and may be stripped intact from the other tissues. Development of the embryonic sporophyte is very similar to that in Desmarestia, as is the anatomy of the adult thallus and the sporangia. From these considerations, Himantothallus is assigned to the Desmarestiaceae (Desmarestiales).     

SPORE WALL ULTRASTRUCTURE OF PROTOSALVINIA

Protosalvinia is an enigmatic fossil which has been historically assigned to several major taxonomic groups. Stratigraphically, the fossil occurs in a narrow range of Upper Devonian sediments. Tetrads of spores are associated with shallow depressions on the surface of approximately 5% of the specimens collected from the Ohio Shale in Columbus, OH. Spores are approximately 250 μm in diameter and have a spore wall which is composed of at least two distinct layers. The outer layer is coarsely laminated in regions where adjacent spores are in contact. Individual laminar units are thinnest toward the inside and gradually thicker toward the surface of the spore. In non-contact regions, the outer layer is composed of globular units. The inner layer of the wall has little discernable structure except for the presence of a distinct suture beneath the proximal trilete mark. This firmly establishes the meiotic nature of these structures. Comparison with eggs and tetraspores of several extant phaeophycean algae shows little similarity.     

THE IDENTITY AND REPRODUCTIVE STRUCTURES OF A MISPLACED SOLENOPORA (RHODOPHYCOPHYTA) FROM THE ORDOVICIAN OF SOUTHWESTERN OHIO AND EASTERN INDIANA1

A study of certain fossil, limestone-forming algae in upper Ordovician beds (Elkhorn and Whitewater Formations of the Richmond Subseries of the Cincinnati Series) in southwestern Ohio and adjacent eastern Indiana culminated with the conclusion that an organism originally described as a sponge, and later placed in the genus Girvanella Nicholson and Etheridge (Cyanochloronta), should in fact be transferred to the genus Solenopora Dybowski (Rhodophycophyta), A calcified reproductive layer was discovered in one of the specimens sectioned. These structures are interpreted as either sporangia or as sporangial chambers, and occur in a sorus rather than a conceptacle type of arrangement. This find has an important bearing on the phylogenetic history of the Solenoporaceae.     

Identification of true Gloiopeltis furcata (Gigartinales,Rhodophyta) and preliminary analysis of its biogeography

In the red algal genus Gloiopeltis, five species (G. complanata, G. dura, G. frutex, G. furcata and G. tenax) are currently recognized, but genetic analyses have suggested considerably greater species diversity. Gloiopeltis specimens formed nine highly supported clades, six of which are morphologically referable to G. furcata. In order to identify the clade corresponding to true G. furcata, we examined the morphology and genetics of the type specimen of Dumontia furcata Postels & Ruprecht, the basionym of G. furcata, housed in the Herbarium of V.L. Komarov Botanical Institute (LE). A clade comprising the type and specimens from the northwestern to northeastern Pacific Ocean was determined to correspond G. furcata. We concluded that the type locality of G. furcata is Sitka, Alaska, based on the identity of haplotypes. We investigated the genetic divergence of populations within this species, using specimens covering most of its distributional range. We confirmed the merger of G. minuta and G. dura with G. furcata.     

ERUMPENT CEPHALODIA,AN APPARENT CASE OF PHYCOBIAL INFLUENCE ON LICHEN MORPHOLOGY1,2,3

On the upper surface of some specimens of the lichen Lobaria cf. crosa (Eschw.) Nyl. there are shrub-like growths which have a morphology similar to that of a free-living fruticose lichen, Polychidium umhausense (Auersw.) Henss., and contain a cyanophyte phycobiont, Nostoc. A few growths also contain scattered colonies of a chlorophyte phycobiont, in which case the lichen tissue locally assumes the foliose form characteristic of the parent Lobaria thallus. The differentiation of dorsiventral lichen cortices and the formation of a lax medulla and distinct algal layer are correlated with the presence of the green phycobiont. The lichen substances atranorin, gyrophoric acid, and 4-0-methylgyrophoric acid occur in both, the foliose L. erosa thallus and the fruticose tissue. It is suggested that the fruticose structures are erumpent cephalodia which are derived from the outgrowth of internal, cephalodia and should not be considered, to be epiphytic colonies of the lichen Polychidium.     

THE RHIZOMORPH APEX OF PAURODENDRON; IMPLICATIONS FOR HOMOLOGIES AMONG THE ROOTING ORGANS OF LYCOPSIDA

A rhizomorph of Paurodendron with an intact apex recently has been discovered in Upper Pennsylvanian sediments of Ohio, and this provides the anatomical evidence necessary to interpret structure, ontogeny and homologies among lycophyte rooting organs. The basal meristem of Paurodendron is radial and lenticular, and produces an apical plug of parenchymatous tissue similar to a root cap. The plug is surrounded by a furrow associated with radially aligned cells that demonstrate a developmental correspondence to the furrow(s) of Isoetes. Based on external structural similarities at the rhizomorph apices of Paurodendron, Stigmaria, and young Nathorstiana, and on the anatomical similarities of Paurodendron to Isoetes, Stigmaria, Chaloneria, and Lepidocarpon embryos, all are interpreted as having a rooting organ that represents a modified shoot system that is fundamentally unlike the primary root system of seed plants. Likewise, the rootlets of rhizomorphic lycophytes are interpreted as leaves modified for rooting, and that have the equivalent of exogenous origin. As such, they are fundamentally unlike the adventitious roots of rhizomatous lycophytes like Lycopodium and Selaginella.     

Euopsis and Harpidium,Genera of the Lichinaceae (Lichenes) with Rostrate Asci*

Based on corresponding ascocarp ontogeny and thallus structure, the genera Euopsis and Harpidium are included in the family Lichinaceae. In the two species of Euopsis, E. granatina and E. pulvinata, the apothecia develop from ascogonia in generative tissue, while in H. rutilans they are pycnoascocarps. In thallus anatomy, the species of Euopsis resemble Pyrenopsis haematopsis and allied species, while H. rutilans corresponds in structure and development of the thallus and apothecia to Pyrenopsis haemaleella (syn. P. sphinctotricha). H. rutilans is the first member of Lichinaceae known to have only a green algal symbiont. In E. granatina, two phycobionts are always present, a species of Gloeocapsa and a chlorococcalean alga. In Euopsis and Harpidium, the ascus wall is composed of an outer, non-expansible and an inner, expansible layer; the latter surrounds the protoplast as an amyloid collar, which expands during spore release into a long, tapering rostrum. In Euopsis, the outer wall layer is strongly amyloid and the upper part separated from the expanded amyloid rostrum by a non-amyloid zone, appearing like a slit in LM studies. The ultrastructure and function of the ascus in E. granatina has been studied in TEM and is interpreted as functionally unitunicate-rostrate. Unitunicate asci with short rostrum are described for P. haemaleella and P. haematopsis.     

Die parasitische FlechteLecidea insidiosa und ihre Biologie

Lecidea insidiosa, up to now regarded as a lichenicolous fungus without thallus, is a parasitic lichen, growing exclusively onLecanora varia. It kills the plectenchyma of the host, but presumably takes over part of the host-algae to build its own thallus, first inside, then outside of the host thallus. Host and parasite thallus are chemically different.Lecidea insidiosa is not related toLecidea vitellinaria, even though the two were thought to be synonymous for a long time. The species is reported for the first time from the Alps (Steiermark). It is the first known parasitic lichen to occur on a normally wood inhabiting host.

     

SETTLEMENT AND SURVIVAL OF POLYSIPHONIA LANOSA (CERAMIALES) SPORES ON ASCOPHYLLUM NODOSUM AND FUCUS VESICULOSUS (FUCALES)1

The settlement patterns of spores of Polysiphonia lanosa (L.) Tandy on Ascophyllum nodosum (L.) Le Jolis and Fucus vesiculosus L. were studied using a flow tank. Settlement sites were defined as ‘sheltered’ or ‘exposed.’ Surface area calculations revealed non-random settlement on A. nodosum, with higher than expected spore frequencies on the thallus and lateral pits and lower than expected frequencies on the vesicles. Settlement of F. vesiculosus was random and significantly lower than on A. nodosum. On the shore, survival of sporelings from September (post-sporulation) to May (pre-sporulation) was highly non-random on both basiphytes. On A. nodosum, lateral pits ('sheltered') showed the highest survival frequency. Here the proportion of surviving sporelings increased over the study period, whereas the proportion on open thallus area ('exposed') decreased. On F. vesiculosus also preferential survival occurred on ‘sheltered’ sites such as vesicle/thallus interfaces and wounds. Between September and May, all P. lanosa sporelings were lost from ‘exposed’ areas (thallus surface and vesicles). Overall, frequencies of surviving sporelings were much greater on A. nodosum than on F. vesiculosus. These results are discussed with reference to basiphyte morphology, epiphyte removal mechanisms and the survival stratagy of P. lanosa.     

BOTRYOPTERIS FORENSIS (BOTRYOPTERIDACEAE), A TRUNK EPIPHYTE OF THE TREE FERN PSARONIUS

Basal parts of Botryopteris forensis have been discovered rooted within the mantle of the tree fern Psaronius. Specimens occur in Upper Pennsylvanian coal balls from near Steubenville, Ohio, USA. The Botryopteris stems branch profusely, and these shoots are intertwined with the Psaronius roots near the surface of the mantle. They also produce adventitious roots that extend among the Psaronius roots. This material demonstrates that B. forensis was a trunk epiphyte, rather than a rhizomatous terrestrial fern. The B. forensis plant is interpreted to have branched continuously, to ramify, and to maintain itself at the periphery of the growing mantle of Psaronius roots. A new reconstruction of B. forensis is offered showing the large, globose fructifications hanging pendulously from horizontal fronds on emergent shoots. Epiphytes and lianas are common on the trunks of Psaronius, indicating that some Marattiales did not produce leaf skirts.     

Intercellular assimilate translocation in Gracilaria cornea (Gracilariaceae,Rhodophyta)

Assimilate translocation has been identified and characterized in Gracilaria cornea under different conditions. Carbon fixation and translocation were carried out by inserting the base part of the thallus into a bicarbonate labeled solution in seawater and exposing its upper part to the air (open system) or to a non-labeled solution above a rubber septum (closed system). After a pulse-chase treatment in the light, three separate sections of each thallus were extracted by DMF (high moleuclar weight photosynthates) or by ethanol (low molecular weight). The results indicate a high rate of active photosynthate translocation which is directly related to inorganic carbon gradients in the thallus, and probably also to sugar gradients in the thallus. Translocation parameters of Gracilaria cornea are lower than of brown algae, as Gracilaria does not contain specific translocation tissues.     

OBSERVATIONS ON THE HABIT,MORPHOLOGY AND ULTRASTRUCTURE OF CEPHALEUROS PARASITICUS (CHLOROPHYTA) AND A COMPARISON WITH C. VIRESCENS1

Cephaleuros parasiticus Karsten, an endophyte of Magnolia grandiflora L. has been examined with light, and scanning and transmission electron microscopy. The discoid thalli are composed of filaments which ramify throughout the leaf tissues beneath the epidermis. Algal filaments do not penetrate host cells, but do produce black leaf spots which have been mistaken for those caused by the fungus Glomerella cingulate (Ston.) Spauld. and Schrenk. Two distinct thallus types occur, often simultaneously on a single leaf. One bears clusters of zoosporangiate branches which seasonally emerge through the ventral (and rarely, dorsal) surface of the leaf. In contrast, the other thallus type bears gametangia which break through the dorsal leaf surface. Zoosporangia and gametangia have never been found on the same thallus. The zoosporangia are smaller than, but almost identical in shape to, those of C. virescens Kunze. Simple plasmodesmata are present in crosswalls and acetolysis indicates that little or no sporopollenin is present in the cell walls. The ultrastructure of biflagellate gametes and quadriflagellate zoospores is virtually indistinguishable from that reported for C. virescens and similar to that reported for Phycopeltis and Trentepohlia. In both gametes and zoospores there are keeled flagella, overlapping and parallel basal bodies, two 3-layered multilayered structures with microtubular splines, and two medial compound microtubular roots. Pyrenoids, eyespots, flagellar and body scales, striated roots (or rhizoplasts), and distal bands are absent. Two presumptive mating structures are present in each biflagellate gamete, and flagellar collars occur in both types of motile cells. The extreme similarity in motile cell ultrastructure revealed in this interspecific comparison parallels that similarity revealed in intergeneric comparisons.     

CHEMICAL COMPOSITION AND STRUCTURE OF THE CELL WALLS OF THE CONCHOCELIS AND THALLUS PHASES OF PORPHYRA TENERA (RHODOPHYCEAE)1,2

Purified cell walls were prepared from both the conchocelis and thallus phases of Porphyra tenera (Kjellm.). The nitrogen content of cell walls from the conchocelis was significantly greater than that for the thallus cell walls, being 3.35 ± 0.26% and 2.39± 0.03%, respectively. Amino acid analysis revealed important differences. The conchocelis cell wall hydrolyzates were richer in aspartic acid, glutamic acid, methionine, and basic amino acids. The thallus cell wall hydrolyzates, however, contained much more glycine and alanine than did those of the conchocelis. Hydroxyproline was not detected in cell walls of either phase. The neutral sugar content of cell wall hydrolyzates from the thallus was more than double that from the conchocelis being 83.6% and 34.5%, respectively. The former contained predominantly mannose which accounted for 72.2% of the neutral sugars while the latter was principally galactose (49.9%) and glucose (36.4%). Methylation analysis confirmed the presence of cellulose microfibrils in the conchocelis in contrast to xylan microfibrils in the thallus. The results establish that the conchocelis and thallus phases of P. tenera differ markedly in the structure and composition of the cell walls.     

POROSIMETRIC STUDY OF THE LICHEN FAMILY UMBILICARIACEAE: ANATOMICAL INTERPRETATION AND IMPLICATIONS FOR WATER STORAGE CAPACITY OF THE THALLUS

The pore system within the thalli of 13 lichen taxa belonging to the family Umbilicariaceae has been studied by means of mercury intrusion porosimetry. A characteristic bimodal pore size distribution with a central depression around 0.05 μm of equivalent pore radius was obtained in all lichen samples. However, clear differences were found among the pore size distributions of each lichen taxa. The total thallus porosity was undoubtedly related to the anatomy of the medulla. In general, a radial plectenchymatic medulla conferred larger porosity to the thallus than an arachnoidal one. Maximum thallus water content closely depended on the total thallus porosity in the five lichens possessing rhizinomorphs. The species with a similar type of medulla could be grouped together in a multivariate analysis that considered three porosimetric parameters and the maximum thallus water content. Umbilicaria cinereorufescens was the most distinct species, with the lowest values of total porosity and water storage capacity and the largest value of thallus density, apparently due to its scleroplectenchymatic medulla. The pore size distribution existing inside the thallus of the species studied is discussed in relation to the often opposing problems of CO₂ exchange and optimal water relations. Some results pointed to a large influence of the micropores (<0.05 μm) on the water storage capacity of the thallus, while the macropores would have a more important role in gas exchange.     

Chloroplast position and photosynthetic characteristics in two monostromatic species,Monostroma angicava and Protomonostroma undulatum (Ulvophyceae), having a shared ecological niche

Monostroma angicava and Protomonostroma undulatum are monostromatic green benthic algae (Ulvophyceae), which grow together in the same intertidal habitat of Muroran, Hokkaido, Japan, during the spring season. Commonly, both species have a single chloroplast with one pyrenoid per cell. The parietal chloroplast is located on the periphery of the thallus in both species, although the location of the chloroplast differs in the two. In M. angicava , the chloroplast was observed to be arranged on one‐side of the thallus surface, whereas, in P. undulatum , it was dispersed and randomly located on either side of the thallus or on the lateral face. The density of chlorophylls (Chls) assessed from the absorption spectra of the thallus and its solvent extract was higher in M. angicava , which appeared dark‐green in color, than in the light‐green colored P. undulatum . The maximum photosynthetic rate per thallus area (μmol O₂ m^?2 s^?1) was higher in M. angicava , whereas, per total chlorophyll content (μmol O₂ g Chl a + Chl b ^?1 s^?1) was higher in P. undulatum . Both species showed similar efficiency of photosynthesis at light‐limiting conditions. The efficiency of light absorption by photosystem II (PSII ) in P. undulatum was higher than M. angicava , whereas the photoprotective response was higher in M. angicava . This indicates that more energy is utilized in M. angicava to protect its PSII due to the chloroplast position, which has more direct exposure to light and, therefore, lowers the efficiency of light absorption by PSII . The higher density of chlorophylls in M. angicava could explain higher photosynthesis per thallus area, whereas, higher efficiency of light absorption by PSII in P. undulatum could explain higher photosynthesis per total chlorophyll content. The differences in light absorption efficiency and quantum efficiency of PSII might be an important ecological strategy in these two species for their coexistence in the intertidal area.