The spores of Pteris cretica complex (Pteridaceae-Pteridophyta) in America

Spore morphology in the Pteris cretica complex, including Pteris brasiliensis, P. ciliaris, P. cretica, P. denticulata var. denticulata, P. denticulata var. tristicula, P. ensiformis, P. multifida and P. mutilata was examined using light and scanning electron microscope. The spores are trilete, triangular to circular in shape, with an equatorial thickening (= cingulum). Equatorial diameter ranges from 29 to 69 μm, and polar diameter from 18 to 53 μm. Exospore is proximally verrucate and distally rugate, with the exception of Pteris ensiformis, which has cones on both polar faces. Perispore is generally less than 1 μm thick, apparently single-layered in section, and translucent under light microscope. Spheroids were frequently observed on the surfaces of both perispore and exospore. Hyaline spores lacking cingulum were found in all specimens. Pteris brasiliensis, P. cretica and P. denticulata exhibit pronounced polymorphism and, in addition to trilete spores these taxa also produce atypical spores such as tetralete, monolete and intermediate types.     

Abnormal spore morphology and wall ultrastructure in Anemia tomentosa var. anthriscifolia and A. tomentosa var. tomentosa (Anemiaceae)

The spores of Anemia tomentosa var. anthriscifolia and A. tomentosa var. tomentosa were studied focusing the attention on their abnormalities. The study was based on fresh and herbarium material and the spores were examined with light, scanning and transmission electron microscopy. Normal, abnormal and abortive spores were observed in both taxa. The normal spores were trilete, triangular in polar view, and the ornamentation consisted of parallel ridges separated by narrow and smooth grooves. The spores were observed in monads, dyads, triads and tetrads. The abnormal spores were monolete, trilete, tetralete or alete with great variations in size. In fact, some spores were almost double the size of the normal ones. Some differences were also found in the ornamentation of the spores. Aborted and not completely developed spores were also observed in the specimens. The wall ultrastructure of the taxa was studied for the first time. The exospore was two-layered with numerous cavities inside its structure, and the perispore was also two-layered. The results revealed that the sporoderm ultrastructure of both normal and abnormal spores of the taxa analyzed was very similar.     

Some notes on Marattia anglica from the Jurassic of Yorkshire

In this paper the spores of the Jurassic fern Marattia anglica are redescribed. It appeared that at least 30% of the spores have a more or less trilete, instead of the usual monolete, laesura. In Marattia muensteri, M. hoerensis and M. crenulata (three species from the Jurassic), trilete spores also appeared to occur, although in much lower percentages than in M. anglica. The spores of M. anglica are compared with recent Marattia spores and with Jurassic dispersed spores.Furthermore, the systematic position of M. anglica and other Jurassic Marattia species is discussed.     

Late Silurian trilete spores from northern Jiangsu, China

The Late Silurian is generally considered to a particular significant key period in the study of early land vascular plants. A trilete spore assemblage of the Upper Silurian is described from northern Jiangsu, China. This assemblage comprises 11 genera and 20 species of trilete spores (including laevigate, apiculate, perinotrilite, patinate, rarely distally murornate and equatorially crassitate, and three indeterminate trilete miospores forms). It has similarities to those described from coeval assemblages from around the world (e.g., England and South Wales; Tripolitania, Libya; Cornwallis Island, Canadian Arctic; Northwest Spain). The rare cryptospore, only one specimen (Tetrahedraletes sp.) had been found to be associated with the Chinese trilete spore assemblage. The discovery of the trilete spores from Late Silurian rocks indicates the existence of early land plants, some possibly vascular, at that time in northern Jiangsu, China.     

Studies in the Spore Morphology of Vittaria Smith

The spore morphology of 32 species of Vittaria Smith has been described. Spores are ellipsoidal or long ellipsoidal in polar view, and kidney-shaped in equatorial view, (18.2–65) × (39–110.5) × (19.5–52)μ in size, monolete, without perine. The stratification of exine which is generally 1.9–2.6 μ in thickness is distinct or indistinct. When it is distinct, the sexine is thicker than the nexine. The exine is generally orna- mented with indistinct granulose or vague sculpture, and with small verrucae in three species. In some species, there is a striate texture on the surface of the exine. As far as it goes, it is a very interesting phenomenon that has not been mentioned elsewhere in the literature of palynology. According to the references, the spores are trilete in a few species of this genus. But it is only monolete in our investigated meterial. Because the gross-morphology of plant in these genera of Vittariaceae is very similar to one anther, we raise the quastion whether the few species which have trilete spores really belong to this genus or to the other genus of Vittariaceae which have trilete, it may be further considered. On the basis of the spore types, Vittariaceae (E. B. Copeland, 1948) may be divided into two groups the one belongs to monolete spore type, and the other to trilete spore type. Based on morphology and anatomy of the plant, R. C. Ching (1940) separated out some genera which have trilete from Vittariaceae to establish a new family– Antrophyaceae, it is valuable to further exploration.     

A PENNSYLVANIAN CALAMITEAN CONE WITH ELATERITES TRIFERENS SPORES

In 1943 L. R. Wilson described some scattered elater-bearing spores found in a Middle Pennsylvanian coal ball from What Cheer, Iowa, as Elaterites triferens. The spores, averaging 60 μ in diameter, with a trilete scar on the proximal surface and three conspicuous elaters attached to their distal surface, have now been found in a fragment of a cone from the same locality. The cone is similar to Calamostachys in having whorls of sporangiophores, each with four adaxial sporangia, and some secondary xylem at the nodes, but it possibly differs in lacking sterile bracts.     

中国奥陶纪晚期—志留纪早期孢型植物及古地理意义

中国奥陶纪晚期的孢型植物组合以隐孢子为主,不含真正三缝孢,称为Tetrahedraletes medinensis-Dy-adospora murusattenuata-Laevolancis chibrikovae(MMC)组合;中国志留纪早期孢型植物组合以隐孢子为主,包括四分体和单分体,缺失二分体,出现真正三缝孢,并产生不同的类型(光面、具纹饰、具弓形脊等类型),被称为Tet-rahedraletes medinensis-Laevolancis chibrikovae-Ambitisporites avitus/dilutus(MCA)组合。孢型植物化石的某些相似性表明:华南和塔里木板块可能分布于相近的低古纬度,气候特征具有一定的可比性,代表近赤道的热带气候;至少在志留纪早期,华南和印支板块与冈瓦纳大陆距离较近,可能处于同一古气候带,代表相似的生物古地理特征。     

Spore morphology of Serpocaulon A.R.Sm. and related taxa from Brazil (Polypodiaceae)

The epiphytic fern genus Serpocaulon (Polypodiaceae) comprises 42 species nested within the neotropical clade of the family together with genera such as Campyloneurum, Grammitis, Microgramma, Niphidium, Pleopeltis, Pecluma, Polypodium and Terpsichore. Although Serpocaulon is a well-supported genus, its intra-specific relationships are still not completely resolved. The morphological dataset for Serpocaulon and related genera is therefore expanded here with information on spore morphology to provide additional information for phylogenetic consideration. Spores have been collected from herbarium vouchers and acetolysed for light and scanning electron microscopy. All members of the neotropical clade have monolete spores, except for Grammitis and Terpsichore, which have trilete spores. Both verrucate and papillate ornamentation types are recognised. The palynological analysis shows close similarity between spores of Serpocaulon, Campyloneurum, Pleopeltis and Polypodium and supports close phylogenetic relationships between these taxa. Species of the grammitid clade, although nested within the Polypodiaceae, have spore morphology that supports a separate position.     

FTIR characterisation of the chemical composition of Silurian miospores (cryptospores and trilete spores) from Gotland, Sweden

To better understand the biological affinities of cryptospores, micro-FTIR (Fourier transform infrared) spectroscopy analysis has been carried out on isolated specimens from the Upper Silurian of Gotland. The geobiochemical results have been compared to spectra of trilete spores, chitinozoans and leiospheres from the same sample. The palynomorphs are all very well preserved as attested by their pale yellow to orange colour indicative of a low thermal maturity. Micro-FTIR spectroscopy indicates that cryptospores display very similar spectra to those of the trilete spores, which are composed of sporopollenin characterised by absorption bands from aliphatic C-H in methylene (CH₂) and methyl (CH₃) groups, aromatic (C=C and C-H) groups and C=O groups of carboxylic acids. The sporopollenin composition of the cryptospore wall observed here is additional evidence demonstrating the embryophytic affinity of the cryptospores. In addition, several variations in other bands in the spectra of the different miospore morphospecies are evidenced and may be linked to their biological affinity or palaeoecological history.     

Palaeophytogeographical and palaeoecological implications of a miospore assemblage of earliest Devonian (Lochkovian) age from Saudi Arabia

Palynological samples from two boreholes drilled in the eastern part of Saudi Arabia contain a rich assemblage of cryptospores and trilete spores. They are dated as most probably Lochkovian in age. Intriguingly, the cryptospores are very similar to those from a coeval continental palynomorph assemblage from the Old Red Sandstone Continent, whereas the trilete spores show a strong affinity with miospore assemblages from Western Gondwana. It is suggested that cryptospore-producing plants inhabited confined, damp biotopes and had a wide range of climatic tolerance. On the other hand, trilete spore-producing plants inhabited a wider variety of biotopes but were more sensitive to climatic variations. The high degree of similarity between Old Red Sandstone and Saudi Arabian cryptospore assemblages favours palaeogeographic reconstructions where there is close proximity between Western Gondwana and Euramerica and/or a land connection between these palaeoplates. Climatic differences between the Old Red Sandstone Continent and the northern part of Western Gondwana can be invoked to explain the variations in the trilete spore assemblages.     

Sporangia containing Scylaspora from the Lower Devonian of the Welsh Borderland

ABSTRACT. Bulk maceration of Early Devonian (Lochkovian) deposits from the Welsh Borderland has yielded eight specimens of a new type of sporangium characterized by its elongate shape and distinctive spores. The specimens have been examined using scanning electron, transmission electron and light microscopy. The elongate sporangia occur isolated and are fragmented to varying degrees. They contain trilete spores that possess a proximal surface with shallow murornate ornament and a distal surface that is laevigate. The spores belong to the dispersed spore genus Scylaspora , and this is the first report of such spores in situ . Ultrastructural studies demonstrate that the spore walls are bilayered with a lamellate inner layer overlain by an essentially homogeneous outer layer. There is little or no associated extra-exosporal material. To date this is the earliest reported example of lamellate wall ultrastructure in trilete spores. Models of spore wall development are suggested in the light of evidence provided by spore wall ultrastructure. Detailed comparisons of the characters preserved in the fossils (morphological, anatomical and ultrastructural), with those in other fossil and extant plants, currently shed little light on the phylogenetic relationships of these specimens, primarily due to the paucity of comparable data. It is proposed that the plant is probably of vascular status, but in the absence of evidence for vascular tissue, it must be classified as rhyniophytoid.     

Microlepia boluoensis sp. nov. (Dennsteadtiaceae) from Guangdong,China

Microlepia boluoensis Y. Yuan & L. Fu, a new species of the genus Microlepia from Luofu mountain, Boluo county, Guangdong province, China, is described and illustrated. Its gametophytic chromosome number is n =47. Microlepia boluoensis has a similar morphology to M. scyphoformis, M. speluncae and M. hancei, such as creeping rhizomes, bipinnate‐tripinnatified laminas, intramarginal sori, cup‐shaped indusium and trilete spores, but it has a shorter creeping rhizome, larger frond (up to 351 cm), dry chartaceous, longer stipe (up to 139 cm), 23 pairs of primary pinnae, 1–13 sori, and spores with a distinct and raised laesurae.     

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.     

Evidence of non-vascular land plants from the early Silurian (Llandoverian) of Virginia,U.S.A.

Evidence is presented for the earliest known occurrences of non-vascular land plants and of higher, septate fungi. Macerates of carbonaceous silstone lenses from the lower Massanutten Sandstone, early Silurian (Llandoverian) of Virginia, have yielded a diverse assemblage of microfossil elements. Parallel aligned, banded tubes with annular to spiral ribbing and rounded to papilliform ends, membranous cellular sheets, cuticles, trilete spores, small spore tetrads, and septate higher filamentous fungi were recovered from the macerates. The banded tubes are probably a significant analogue with supportive or conductive cell types, but are not considered tracheidal. The heterogeneous plant assemblage may represent a thalloid, non-vascular land plant, in part, with a tubular-filamentous (nematophytic) organization associated with a membranous cellular layer and cuticular covering. While no spores were established as nematophytic, the presence of trilete spores adds to the indirect evidence of multiple evolutionary convergence toward land-plant characters. This assemblage is interpreted as of land-plant origin, based on the inferred fluvial depositional model of the fossiliferous rocks. A glacio-eustatic sea-level drop in the late Ordovician is suggested as a stimulus to the advent of land plants in the early Silurian.     

Lower Devonian coalified sporangia from Shropshire: Salopella Edwards & Richardson and Tortilicaulis Edwards

A diverse assemblage of coalified sporangia from Lochkovian/Gedinnian, fluvial rocks imicrornatus-newportensis Spore Biozone) contains fusiform forms assignable to Salopella Edwards & Richardson and Tortilicaulis Edwards. Both show bifurcation within some sporangia. In specimens assigned to Salopella cf. marcensis , longitudinal dehiscence produces two equal valves revealing spores resembling Aneurospora , although trilete marks may be difficult to distinguish. The spores are in two forms with distal ornament of either coarse or fine coni, each occurring in separate sporangia. A further sporangium of Salopella shape contains distally apiculate, hilate cryptospores. Elongate sporangia exhibiting spiralling of superficial cells and sometimes gross twisting, are placed in a new species of Tortilicaulii, T. offaeus. Anatomical data include details of irregularly thickened epidermal cells, sporangial wall and in situ spores. The latter are trilete, equatorially thickened and highly distinctive, because the entire exospore surface is covered with grana, a feature not common in dispersed spores of this age. The nomenclatural and curatorial problems associated with these small, coalified fossils, whose anatomical examination by scanning electron microscope necessitates elimination of gross morphology, are discussed.     

BOTRYOPTERIS PINNULES WITH ABAXIAL SPORANGIA

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

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.     

TETRAXYLOPTERIS SCHMIDTII: ITS FERTILE PARTS AND ITS RELATIONSHIPS WITHIN THE ANEUROPHYTALES

The fertile branching system of Tetraxylopteris is composed of successive “nodes” bearing opposite and decussately arranged, upcurved sporangial complexes. By means of the transfer technique the morphology of the sporangial complex was revealed. It consists of a main stalk which dichotomizes twice producing four major branches. Each of the four branches is further subdivided three times, the subdivisions being arranged alternately and pinnately. The ultimate divisions bear the sporangia singly and terminally. The sporangial complexes decrease in size distally and are more tightly curved at the apex. The sporangia are oblong-oval with an acute apex. The spores are identical to the dispersed spore taxon Rhabdosporites langii, Richardson. They are spherical, trilete and pseudosaccate with a fine granular ornament on the pseudosaccus. They are 75–176 μ in diameter and show developmental stages from young tetrads to separated, fully mature spores depending on the age of the sporangium from which they were obtained. This is the first account of spores in sporangia of Tetraxylopteris. The diagnosis of the genus and species are emended to include the new information and the order Aneurophytales is redefined.     

CALAMOPHYTON IN THE MIDDLE DEVONIAN OF NEW YORK STATE

Specimens of Calamophyton from the Middle Devonian Ashokan Sandstone near Kingston, Ulster County, N.Y., are shown to belong to C. bicephalum Leclercq and Andrews, a Belgian species. Steel needles and a stereoscopic binocular microscope were used to follow the path of the forking leaves and branching sporangiophores of the specimens through the matrix. The terete leaves dichotomized two to three times in more than one plane. Sporangiophores dichotomized once. Each branch bore three recurved lateral branches which in turn bore two sporangia. Each branch terminated in an elongate, filiform projection. Sporangia apparently dehisced longitudinally. Their walls were composed of elongate cells. Their spherical spores ranged from 86 to 166 μ in diameter and bore a trilete mark. Ornamentation consisted of coni and spinae up to 4.5 μ long. They resembled dispersed spores of Dibolisporites gibberosus var. major Richardson. This is the second occurrence of Calamophyton bicephalum and the first account of its spores. It is the second report of the genus in North America.     

The microfossil record of early land plants

Dispersed microfossils (spores and phytodebris) provide the earliest evidence for land plants. They are first reported from the Llanvirn (Mid-Ordovician). More or less identical assemblages occur from the Llanvirn (Mid-Ordovician) to the late Llandovery (Early Silurian), suggesting a period of relative stasis some 40 Myr in duration. Various lines of evidence suggest that these early dispersed microfossils derive from parent plants that were bryophyte-like if not in fact bryophytes. In the late Llandovery (late Early Silurian) there was a major change in the nature of dispersed spore assemblages as the separated products of dyads (hilate monads) and tetrads (trilete spores) became relatively abundant. The inception of trilete spores probably represents the appearance of vascular plants or their immediate progenitors. A little later in time, in the Wenlock (early Late Silurian), the earliest unequivocal land plant megafossils occur. They are represented by rhyniophytoids. It is only from the Late Silurian onwards that the microfossil/ megafossil record can be integrated and utilized in interpretation of the flora. Dispersed microfossils are preserved in vast numbers, in a variety of environments, and have a reasonable spatial and temporal fossil record. The fossil record of plant megafossils by comparison is poor and biased, with only a dozen or so known pre-Devonian assemblages. In this paper, the early land plant microfossil record, and its interpretation, are reviewed. New discoveries, novel techniques and fresh lines of inquiry are outlined and discussed.