Exine ultrastructure of in situ peltasperm pollen from the Rhaetian of Germany and its implications

Morphology and exine ultrastructure of pollen grains of Triassic peltasperms have been studied for the first time. Pollen grains of Antevsia zeilleri from the Rhaetian of Germany are of the Cycadopites-type and monosulcate; the sculpturing is the same in the apertural and non-apertural areas. The proximal exine includes a row of lacunae covered by a solid, thick tectum and underlined by a foot layer. Pillars are hanging from the tectum between the lacunae. The exine is thinning to a homogeneous layer in the apertural region. The latter is bordered by thicker alveolate areas of the exine, in places resembling a saccus-like ultrastructure. The endexine includes white-line-centred lamellae. The exine ultrastructure is compared with that of pollen of Permian peltasperms. Although pollen types ascribed to Permian peltasperms are completely different in their general morphology, a transformation can be hypothesized by ultrastructural data from Permian Vesicaspora into Triassic Cycadopites extracted from pollen sacs of Antevsia. Comparison with Cycadopites of non-peltaspermalean (Ginkgoalean, Cycadophyte) and unknown affinities has been accomplished. The exine ultrastructure is distinctive enough to differentiate among peltaspermalean, cycadalean and bennettitalean Cycadopites; some ultrastructural features are shared with pollen of modern Ginkgo biloba. More ultrastructural data are needed as well as numerous sections of pollen grains are necessary to reveal original unchanged ultrastructure.     

Exine micromorphology and ultrastructure in Neottieae (Epidendroideae,Orchidaceae)

The diverse epidendroid orchid tribe Neottieae is characterized by multiple transitions between autotrophy and mycoheterotrophy, allogamous and autogamous mating systems, pollen released as tetrads or monads, and pollen exine tectate or semitectate. We use transmission and scanning electron microscopy on pollen of ten species of Neottieae to investigate whether the differences in pollen aggregation and exine micromorphology and ultrastructure reflect phylogenetic relationships, or whether this variation is subject to ecological constraints. Our results showed that differences in exine micromorphology are mostly concordant with phylogenetic relationships in Neottieae, i.e. an ascending tendency of pollen ornamentation from tectate (Cephalanthera) to semitectate (e.g. Neottia). In contrast, pollen aggregation, when plotted on the most recent phylogeny, shows repeated transitions between monads and tetrads that could be related to ecological constraints. Tetrads are present in species that are nectar rewarding, whereas monads are common in deceptive species. Cephalanthera is characterized by recalcitrant pollen, including the frequent occurrence of collapsed pollen. In this genus, the observed shifts from allogamous to autogamous or cleistogamous mating systems could help to reduce pollen damage caused by exposure to dry habitats.     

Rhaetian magneto-biostratigraphy from the Southern Alps (Italy): Constraints on Triassic chronology

New Late Triassic–earliest Jurassic magneto-biostratigraphic data have been obtained from three overlapping sections in the Southern Alps, Italy (Costa Imagna, Brumano, Italcementi Quarry), composed of ~ 520 m of shallow-marine carbonates outcropping in stratigraphic continuity. Characteristic magnetic components of presumed depositional age record a sequence of 9 normal and reverse polarity magnetozones (as defined by at least three stratigraphically superposed samples) linked by conodont and palynofloral evidence from this study and the literature to Rhaetian to Triassic–Jurassic boundary age. This represents a significantly larger number of polarity zones than previously recognized in more condensed Rhaetian sections from the literature, and by inference represents more time. These data are placed in a broader Late Triassic temporal framework by means of correlations to published magneto-biostratigraphic data from the Tethyan marine Pizzo Mondello section and the Newark astronomical polarity time scale (APTS). This framework is consistent with a position of the Norian–Rhaetian boundary (as defined at Brumano and Pizzo Mondello by the first appearance of Misikella posthernsteini) within Newark magnetozones E17r–E19r in the ~ 207–210 Ma time interval, in basic agreement with the position originally estimated in the Newark using pollen biostratigraphy (E18 at 208–209 Ma). This framework is also consistent with the position of the Triassic–Jurassic boundary interval (placed at Italcementi Quarry at the acme of Kraeuselisporites reissingeri coincident with a negative carbon isotope excursion) correlative to just above Newark magnetozone E23r and just below the oldest CAMP lavas dated at ~ 202 Ma. Hence, we estimate the duration of the Rhaetian to be ~ 5.5–8.5 Myr (or even longer if the Triassic–Jurassic boundary is instead placed above the negative carbon isotope excursion as at Kuhjoch, which is the designated GSSP for the base of the Hettangian), and encompassing 9 magnetozones. This duration contrasts with a duration of ~ 2 Myr and only ~ 4 magnetozones in several alternative published magneto-biostratigraphic schemes.     

A Reappraisal of the Purported Gastric Pellet with Pterosaurian Bones from the Upper Triassic of Italy

A small accumulation of bones from the Norian (Upper Triassic) of the Seazza Brook Valley (Carnic Prealps, Northern Italy) was originally (1989) identified as a gastric pellet made of pterosaur skeletal elements. The specimen has been reported in literature as one of the very few cases of gastric ejecta containing pterosaur bones since then. However, the detailed analysis of the bones preserved in the pellet, their study by X-ray microCT, and the comparison with those of basal pterosaurs do not support a referral to the Pterosauria. Comparison with the osteology of a large sample of Middle-Late Triassic reptiles shows some affinity with the protorosaurians, mainly with Langobardisaurus pandolfii that was found in the same formation as the pellet. However, differences with this species suggest that the bones belong to a similar but distinct taxon. The interpretation as a gastric pellet is confirmed.     

Sexual dimorphism in a new species of the actinopterygian Peltopleurus from the Triassic of northern Italy

The new species Peltopleurus nuptialis is described on the basis of several well preserved specimens from the uppermost Ladinian Kalkschieferzone of Ca' del Frate. This new species is characterized by small size and the presence of hook-like fulcra on all fins except the caudal, and by tubercles on the rostral and nasal bones interpreted as secondary sexual traits. There is also a modified anal fin in supposed males, which probably acted as a gonopodium, pointing to a strong sexual dimorphism. A distinctive morphological variability seems to characterize peltopleurids: a comparison with groups of living fishes also showing morphological plasticity and sexual dimorphism is given in order to hypothesize similar behaviours and adaptations in the fossil forms.     

A Tetrapod Ichnoassociation from the Middle Triassic (Anisian,Pelsonian) of Northern Italy

A new Middle Triassic ichnofauna recovered nearby the town of Tisens (Bozen) in the Adige Valley (Trentino, Northern Italy) is described. A relatively large ichnoassociation, in which various ichnogenera can be recognized, comes from the basal inter-supratidal layers of a mixed carbonate siliciclastic unit of Pelsonian (Middle Anisian) age. Most of the isolated footprints and trackways pertain to lizard-like reptiles referable to Rhyncosauroides and, subordinately, to archosaurian reptiles. In particular, the ichnogenera Rotodactylus, Synaptichnium and Chirotherium have been recognized. Many tracks are at present unidentified; among them a lizard-like morphotype (morphotype A). The site also yelded numerous fossil plant horizons and some scattered vertebrate remains referable to Placodontidae (cf. Placodus gigas). The site shows the continuity in the Pelsonian of the “chirothere ichnofauna” and the dominance of Rhynchosauroides in carbonate, tidal flat environments.     

A new dasycladalean alga from the Middle Triassic carbonates of Northern Calabria (Southern Italy)

Summary A new dasyclad alga—Acicularia boniae n.sp.—is discribed from Middle Triassic (?Ladinian) of the Piano del Minatore Formation, outcropping in the Cozzo del Pellegrino area (Calabria, southern Italy). The systematic position of this species, with respect to the genusAcicularia and morphogeneraAciculella andTerquemella, is discussed.

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Riassunto Viene descritta una nuova specie di dasicladale—Acicularia boniae—proveniente dal Triassico Medio della formazione del Piano del Minatore affiorante nel massiccio del Cozzo del Pellegrino (Calabria, Italia meridionale). Viene discussa la posizione sistematica rispetto al genereAcicularia ed ai morfogeneriTerquemella edAciculella.

     

Exine ultrastructure in pollen grains of <Emphasis Type=Italic>Classopollis</Emphasis> Pflug from the Cretaceous of Lebanon

Pollen grains of Classopollis Pflug from the Cretaceous deposits of Lebanon were studied by means of light and electron microscopy. Ultrastructurally, they are similar to pollen grains, extracted from Classostrobus comptonensis Alvin, Spicer et Watson from the Barremian of England. The differences concern the shape and size of spinules, ultrastructure of apertural regions, and preservation of the endexine. An analysis of our data and published results revealed three types of infratectum existed in members of Circumpolles: (1) with branchy elements, (2) with columella-like non-branching elements, and (3) with large granules arranged in one row. The palynological assemblage is described in detail; problems of dating are discussed.     

Characteristics of morphology and ultrastructure of bacteria of the genus Dethiosulfovibrio

Cell morphology and fine structure were studied in two strains of rod-shaped, strictly anaerobic, gram-negative sulfidogenic bacteria: strain SR12T (DSM 12538) and strain WS100 (DSM 12537) belonging to "Dethiosulfovibrio starorussensis." Cells of both strains, as well as cells of the type species of the genus Dethiosulfovibrio, D. peptidovorans, were found to possess multiple intracellular incomplete cross septa in the stationary growth phase.     

Pollen morphology and ultrastructure of selected species of Magnoliaceae

The pollen morphology and ultrastructure of 20 species, representing eight genera of the Magnoliaceae are described based on observations with light, scanning and transmission electron microscopy. The family represents a homogeneous group from a pollen morphological point of view. The pollen grains are boat-shaped with a single elongate aperture on the distal face. The tectum is usually microperforate, rarely slightly or coarsely rugulose. Columellae are often irregular, but well-developed columellae do occur in some taxa. The endexine is distinct in 14 species, but difficult to discern in the genera Parakmeria, Kmeria and Tsoongiodendron. Within the aperture zone the exine elements are reduced to a thin foot layer. The intine has three layers with many vesicular-fibrillar components and tubular extensions in intine 1. The symmetry of the pollen grains, shape, type of aperture and ultrastructure of the intine show a remarkable uniformity in the family. Nevertheless there is variety in pollen size, ornamentation and the ultrastructure of the exine. The pollen of Magnoliaceae is an example of an early trend of specialization, and supports the view that Magnoliaceae are not one of the earliest lines in the phylogeny of flowering plants.     

Spore morphology and wall ultrastructure of Hymenophyllaceae Link (Pteridophyta) from north-west Argentina

The family Hymenophyllaceae is represented in the study area by six species in two genera, Hymenophyllum J. E. Smith and Trichomanes L. The study was based on herbarium material and spores were studied under light microscope (LM), scanning electron microscope (SEM) and transmission electron microscope (TEM). Both genera have trilete spores, 23 to 45 μm in equatorial diameter, with an ornamentation of echinulae and cones in Hymenophyllum and of verrucae, gemmae and granules in Trichomanes. Mature spores have a sporoderm composed of a perispore, an exospore and a fibrillar endospore; the exospore is 0.5 to 2.5 μm thick, compact and with an irregular margin. In some cases radial channels and other channels associated with the middle and inner parts of the laesurae were evident. A series of cavities filled with an opaque content line the inner margin of the exospore. The perispore is 20 to 400 nm thick and unevenly differentiated along the surface of a same spore. Under TEM, two main differentially contrasted portions could be distinguished: a dark massive portion with structural components could not be distinguished, and a light portion with several plates arranged in piles. The inner surface of the perispore exhibit short scales. Globules are immersed within the perispore at some depth from the perispore surface and others connected to it by structural threads. The spore characters observed including shape, ornamentation, laesurae length and wall structure are useful in distinguishing the two genera studied, but less useful in differentiation at the species level.     

On the morphology and ultrastructure of the cell wall of Spirulina platensis

The cell wall of the blue-green alga Spirulina platensis was studied with the electron microscope using ultra-thin sectioning, shadowing, carbon-replication or freeze-etching techniques for specimen preparation. The cell wall could be resolved into four layers, L-I through L-IV. The L-I and L-III layers contain fibrillar material. The septum is a three-layered wall: an L-II layer sandwiched between L-I layers. The shape in vitro of isolated septa might be an artifact due to the preparation technique used. Certain structural properties of the septum seem to allow tangential stretching; they might be reflected in the flexible gliding mobility of Spirulina species. The outer, L-IV layer contains material longitudinally arranged along the trichome axis.     

Exine and tapetum development in Symphytum officinale (Boraginaceae). Exine substructure and its interpretation

After detailing the exine ontogeny, our purpose was to find out whether the sequence of sporoderm developmental events corresponds to self-assembling micellar mesophases, initiated by genomically determined physicochemical parameters and induced by surfactant glycoproteins at increasing concentrations. Indeed, a scaffolding of the future exine, i.e., the glycocalyx, initiates with scattered clots, which then appear as clusters of spherical and worm-like micelles, derived from surface-active glycoproteins. At the middle tetrad stage, a continuous layer of the glycocalyx emerges, consisting of parallel, tightly packed cylinder-like units, which we interpret as a layer of cylindrical micelles, the so-called middle mesophase. These units bear dark-contrasted particles, arranged in strings or columns. These sites of the glycocalyx units?Cmicelles accumulate initial sporopollenin, hence the term ??sporopollenin acceptor particles?? (SAPs). This process leads to the appearance of procolumellae at the late tetrad stage. The glycocalyx units are rooted into callose and into the microspore cytoplasm. After formation of the tectum and the foot layer, the endexine initiates as a thin layer, and the latter develops into a very thick layer in the post-tetrad period. When callose disintegrates, ??bouquets?? of SAPs become evident on the tectum, which were evidently hidden inside the callose layer; these structures self-assemble into supratectal gemmae. An unusual, ??hybrid?? type of tapetum was observed. What is observed in Symphytum exine development allows us to obtain more evidence for the hypothesis of the participation of micellar self-assembly in sporoderm development and to bring together the concepts of micelles and of SAPs.     

Floral Assemblages and Patterns of Insect Herbivory during the Permian to Triassic of Northeastern Italy

To discern the effect of the end-Permian (P-Tr) ecological crisis on land, interactions between plants and their insect herbivores were examined for four time intervals containing ten major floras from the Dolomites of northeastern Italy during a Permian–Triassic interval. These floras are: (i) the Kungurian Tregiovo Flora; (ii) the Wuchiapingian Bletterbach Flora; (iii) three Anisian floras; and (iv) five Ladinian floras. Derived plant–insect interactional data is based on 4242 plant specimens (1995 Permian, 2247 Triassic) allocated to 86 fossil taxa (32 Permian, 56 Triassic), representing lycophytes, sphenophytes, pteridophytes, pteridosperms, ginkgophytes, cycadophytes and coniferophytes from 37 million-year interval (23 m.yr. Permian, 14 m.yr. Triassic). Major Kungurian herbivorized plants were unaffiliated taxa and pteridosperms; later during the Wuchiapingian cycadophytes were predominantly consumed. For the Anisian, pteridosperms and cycadophytes were preferentially consumed, and subordinately pteridophytes, lycophytes and conifers. Ladinian herbivores overwhelming targeted pteridosperms and subordinately cycadophytes and conifers. Throughout the interval the percentage of insect-damaged leaves in bulk floras, as a proportion of total leaves examined, varied from 3.6% for the Kungurian (N = 464 leaves), 1.95% for the Wuchiapingian (N = 1531), 11.65% for the pooled Anisian (N = 1324), to 10.72% for the pooled Ladinian (N = 923), documenting an overall herbivory rise. The percentage of generalized consumption, equivalent to external foliage feeding, consistently exceeded the level of specialized consumption from internal feeding. Generalized damage ranged from 73.6% (Kungurian) of all feeding damage, to 79% (Wuchiapingian), 65.5% (pooled Anisian) and 73.2% (pooled Ladinian). Generalized-to-specialized ratios show minimal change through the interval, although herbivore component community structure (herbivore species feeding on a single plant-host species) increasingly was partitioned from Wuchiapingian to Ladinian. The Paleozoic plant with the richest herbivore component community, the coniferophyte Pseudovoltzia liebeana, harbored four damage types (DTs), whereas its Triassic parallel, the pteridosperm Scytophyllum bergeri housed 11 DTs, almost four times that of P. liebeana. Although generalized DTs of P. liebeana were similar to S. bergeri, there was expansion of Triassic specialized feeding types, including leaf mining. Permian–Triassic generalized herbivory remained relatively constant, but specialized herbivores more finely partitioned plant-host tissues via new feeding modes, especially in the Anisian. Insect-damaged leaf percentages for Dolomites Kungurian and Wuchiapingian floras were similar to those of lower Permian, north-central Texas, but only one-third that of southeastern Brazil. Global herbivore patterns for Early Triassic plant–insect interactions remain unknown.     

Pollen morphology and ultrastructure of Marathrum schiedeanum (Podostemaceae)

The Podostemaceae, or river-weeds, comprise 46 genera and 270 species of dicots and are the largest family of strictly aquatic angiosperms. Despite the large size, specialized habitats, and enigmatic morphology of the family, relatively little is known about the palynology of Podostemaceae. In the current paper, pollen morphology and ultrastructure of Marathrum schiedeanum are described. Pollen grains are relatively small, spheroidal, and tricolpate to spiraperturate. The exine has a microechinate ornament, a tectate-granular sexine and a relatively thick nexine in non-apertural regions, and a semitectate sexine and thinner nexine in apertural regions. Although aperture variation occurs in the family, this is the first report of the spiral aperture type in Podostemaceae. The spiraperturate condition appears to be derived in river-weeds, as does the granular pollen wall, which represents a reduction of the typical columellae found in eudicots.     

Topography and Non-Topography in Exine Organization

Table I shows that two letters (S, N) and two figures (1, 2) can be used as symbols or “labels” instead of more or less complicated and/or ambiguous terms in labelling the main topographical and/or non-topographical sub-units or parts of angiosperm exine types (at least as far as present knowledge goes). Special attention has been paid to the characteristics of the nexine in Epilobium montanum believed to be “nexine-2” (N2) by Erdtman and others, albeit classified as “nexine-1, footlayer of the ektexine” by Rowley et al. (1970; p. 7).