Late Silurian plant microfossil assemblage from Guangyuan,Sichuan, China

A dispersed plant microfossil assemblage is described from Late Silurian deposits from Guangyuan, Sichuan, China. These strata are interpreted as nearshore, shallow marine deposits, and brachiopods suggest a late Ludlow–early Pridoli age. The palynomorph assemblage is dominated by terrestrial forms, including cryptospores and trilete spores, tubular structures and cuticle-like sheets, although rare marine acritarchs are also present. This microfossil assemblage is comparable to coeval assemblages from around the world (South and Southwest Wales; Libya; Canada; Southeastern Turkey; Northwest Spain; and Jiangsu, China). The sporomorphs from this assemblage indicate the existence of early land plants during the late Ludlow–early Pridoli in Guangyuan, Sichuan, China; and suggests that floras of this age were cosmopolitan and exhibited little palaeogeographical differentiation.     

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.     

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.     

Early vascular land plants: proof and conjecture

Megafossil evidence does not fill the 'evolutionary gap' between land plants and their hypothetical green algal ancestors. Rare Late Silurian vascular plant megafossils provide little information about the morphological, physiological, biochemical, and ecological steps that preceded their evolution. Dissociated trilete spores, spore tetrads, cuticle- and tracheid-like structures far exceed the abundance and diversity of Silurian vascular plant megafossils, and appear millions of years before them. In reference to whole-bodied organisms, these or analogous structures belong to land plants or emergent aquatics; they may represent plants evolutionarily intermediate between green algae and descendent vascular plants at the bryophyte or pre-bryophyte stages. Changes in the cellular biochemistry of pre-Devonian land plants in response to the selective pressures of terrestrial life may have led to the origin of lignin and cutin, neither of which has any counterpart among the algae, and to the evolutionary surge of the vascular plants in the Early Devonian represented by the plant megafossil record. Positive correlation between abundance and diversity of trilete spores and shallow-water, nearshore sites reinforces conclusions based on morphology that a terrestrial flora existed well prior to the appearance of vascular plant megafossils.     

Implications of an early land plant spore assemblage for the late Silurian age of the Si Ka Formation,northern Vietnam

The first plant microfossil assemblage from the Si Ka Formation of the Song Cau Group, northern Vietnam is reported. It is composed of cryptospores in dyads and tetrads, trilete spores, tubular remains consisting of an association of smooth, banded, and externally thickened tubes, and cuticle-like fragments. The biostratigraphic assemblage of sporomorphs indicates a late Silurian (late Ludfordian) to Early Devonian (early Lochkovian) age. Further comparison with coeval reports using the characteristic features of the assemblage confines their age to the late Ludlow (late Ludfordian) to early Přídolí. This report presents the oldest spore assemblage from Vietnam and contributes to a broader understanding of its paleo-landscape during the late Silurian.     

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。     

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.     

Devonian Spores in a Middle Silurian Reef

A subsurface core from the Tilden reef (Middle Silurian) in southern Illinois contains a large and diverse assemblage of hystrichospherids and plant spores. The plant spores are restricted to sinuous cracks filled with calcite and brownish-gray clay. The miospore assemblage is dated late Middle or early Late Devonian, and thus the fissure fillings represent a “stratigraphic leak”. Failure to recognize this phenomenon could result in misleading age determinations.

The spore assemblage is dominated by cingulate miospores of the genera Stenozonotriletes and Retusotriletes and by spores with perisporal membranes assignable to Diaphanospora. Seventeen new species are described and illustrated: one each of Deltoidospora, Anapiculatisporites, Verrucosisporites, Cristatisporites, Rhabdosporites, Ancyrospora, and Endosporites; two each of Converrucosisporites and Retusotriletes, and three each of Stenozonotriletes and Diaphanospora.     

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.     

Spore assemblages from the Lower Devonian Xujiachong Formation from Qujing,Yunnan, China

Abstract: The Lower Devonian Xujiachong Formation from the vicinity of Qujing City, Yunnan, China is interpreted as a terrestrial‐fluviatile‐lacustrine sequence. It contains important nonmarine biotas including plants, fish and invertebrates. The plants are particularly interesting as they include many endemic taxa. Dispersed spore assemblages have been recovered from the upper part of this formation. The spores are well preserved and of moderate thermal maturity. They are systematically described and four new species erected: Aneurospora xujiachongensis sp. nov., Chelinospora ouyangii sp. nov., Camptozonotriletes? luii sp. nov. and Leiozonospora xichongensis sp. nov. One new combination is proposed: Aneurospora conica (Ouyang and Lu) comb. nov. This is a rare report of a Lower Devonian dispersed spore assemblage from the South China Plate. Indeed, few dispersed spore assemblages of this age are known outside of Euramerica and Northern Gondwana. It is suggested that the Xujiachong Formation spore assemblages can all be equated to the polygonalis‐emsiensis Spore Assemblages Biozone (PE SAB) of Richardson and McGregor (1986) indicating an early (but not earliest) Pragian to ?earliest Emsian age. However, caution is urged, because biostratigraphical interpretation is difficult owing to distinct differences between dispersed spore assemblages from South China and Euramerica/Northern Gondwana. This almost certainly reflects palaeophytogeographical variation and regional endemism among early land plant floras on widely separated land masses. Palynofacies analysis supports a nonmarine origin for the deposits of the Xujiachong Formation, with the very rare marine palynomorphs that were encountered interpreted as reworked.     

新疆若羌石灰华中的植物化石及古环境*

根据首次从富阳西湖组发现的孢子及其垂直分布特征,划分出下段 LH 组合和上段 DP 组合,分别归入晚泥盆世法门晚期(斯图年期)和早石炭世早期(杜内早期),即泥盆-石炭系界线通过西湖组内部,与江苏五通组类似.     

Ultrastructure of laevigate hilate spores in sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland

Spore masses and isolated sporangia, containing laevigate hilate cryptospores attributable to the dispersed taxon Laevolancis divellomedia sensu lato, have been recovered on bulk maceration of Upper Silurian (Pridoli) and Lower Devonian (Lochkovian) deposits from the Welsh Borderland. Detailed morphological, anatomical and ultrastructural analysis, using light microscope, scanning electron microscope and transmission electron microscope techniques, reveals subtle differences between the specimens and they can be grouped into five distinct types. The different groups are distinguished principally by using sporangia-spore mass characteristics, presence or absence of extra-exosporal material and nature of spore-wall ultrastructure. Of the groups, one has a uniformly homogeneous exospore and the other four groups have a bilayered exospore. In the former the spores lack extra-exosporal material and occur in a discoidal sporangium. Of the bilayered groups, two have exospores of homogeneous composition but with the two layers differing in electron density. They occur in discoidal sporangia and spore masses and are distinguished on the presence or absence of extra-exosporal material and differences in the widths of the two layers. Finally, two bilayered groups possess a lamellate inner layer, but vary in presumed sporangial shape. Elongate sporangia have spores with concentric continuous lamellae, lacking further ultrastructure. In contrast, spores from a discoidal spore mass have white-line-centred, presumably tripartite, lamellae which are laterally discontinuous, overlapping and irregularly spaced. These findings, which suggest that morphologically similar spores were produced by a number of plant taxa, have important implications regarding the assessment of early land-plant diversity. The affinities of hilate cryptospore-producing plants are unknown and problematic, particularly as no extant non-angiosperm plants produce dyads, other than through meiotic irregularity, and spore-sporangial characters have no exact counterpart in coeval plants. Studies of specimens with in situ hilate cryptospores suggest that they derive from rhyniophytoids, i.e. plants that resemble the simplest of vascular plants but lack evidence of vascular tissue, although hilate cryptospore-containing examples show no axial branching. It might be argued, based on evidence from spore wall ultrastructure, that some of the plants have more in common with lycopsids and filicopsids than bryophytes, a surprising finding bearing in mind the stratigraphic distribution of hilate cryptospores-dyads and inferences that the producers were bryophyte-like. Detailed studies of wall structure in the hilate cryptospores permit consideration of spore wall development. It is suggested that extra-exosporal material derives from a tapetum and is thus produced by the diploid sporophyte. The white-line-centred lamellae in a single specimen provide the earliest evidence for the presence of such structures in early land plant spores and provide further evidence that sporopollenin deposition on such structures is the most primitive mode of sporopollenin deposition among land plants.     

Llandovery sporomorphs and graptolites from the Manbo Formation,the Mojiang County,Yunnan, China

Llandovery sporomorphs and graptolites have been recovered from the Manbo Formation, Mojiang area, western Yunnan, China, which belonged to the Indo-China Palaeoplate in the Palaeozoic. The graptolite fauna, including 8 genera and 12 species, is considered Mid Telychian, Llandovery in geological age, confirming the existence of Llandovery rocks in this region. The Silurian stratigraphical sequence of this area is reconsidered as in the ascending order: Manbo Formation (Llandovery–early Wenlock), Shuiqing Formation (late Wenlock–Ludlow) and unnamed formation (Pridoli?). The sporomorph assemblage from the Manbo Formation includes seven species in five genera. The dominant members of this assemblage are Tetrahedraletes medinensis and Laevolancis chibrikovae. Based on the low diversity and low abundance, the geological age of this assemblage is considered to be approximately the Telychian, Llandovery. The parent plants of sporomorphs probably inhabited the landmass near Mojiang area, these sporomorph-producing plants probably included bryophyte-like land plants and primitive vascular land plants. The sporomorph evidence shows that the South China and Indo-China palaeoplates may have been in close proximity (maybe with some continental bridges linking them) to each other at least in the Llandovery. They were also closely related with Gondwanaland in the Llandovery.     

The development of early terrestrial ecosystems

Summary

In this review of terrestrialization by plants and animals in the early Phanerozoic, the classical idea of a major mid-Palaeozoic event is discarded in favour of gradual colonization over a long time period. Four phases of colonization of the land by plants are recognized but their limits are often difficult to define. The first, of microbial mats comprising prokaryotes and later photosynthesizing protists (algae) but with no direct fossil evidence, extends from the Precambrian and may persist in environments unsuitable for colonization by higher plants and animals today. The second, based on microfossils (spores and cuticles) possibly from plants of bryophyte aspect (if not affinity) started in the Ordovician (c. 460 Ma ago) and ended in the Lower Devonian, but was overlapped by the third phase beginning early in the Silurian (c. 430 Ma). This consisted of small plants of axial organization with terminal sporangia probably allied to the tracheophytes. The advent of taller vascular plants of varied organization around the Silurian — Devonian boundary (c. 420–400 Ma) signalled the final pioneering phase — that of major adaptative radiations on land, culminating in the appearance of extant groups, in changes in reproductive strategy and in the development of complex vegetation structure. The animal record is sparser than that of the plants, but suggests an early land fauna in the mid-Palaeozoic which differed from later terrestrial assemblages in lacking herbivores, with the first direct fossil evidence for land animals in the late Silurian.     

Composition and dynamics of the great Phanerozoic Evolutionary Floras

Factor analysis of a data set representing the global distribution of vascular plant families through time shows the broad pattern of vegetation history can be explained in terms of five Evolutionary Floras. The Rhyniophytic (=Eotrachyophytic) Flora represents the very earliest (Silurian and earliest Devonian) vascular plants, notably the Rhyniophytopsida. The Eophytic Flora represents the early (Early–Middle Devonian) mainly homosporous land plants, notably the Zosterophyllopsida, Trimerophytopsida and early Lycopsida. The Palaeophytic Flora represents the Late Devonian and Carboniferous vegetation, which saw the introduction of heterospory among the spore producing plants and of early gymnosperms. The Mesophytic Flora first appeared in the Late Carboniferous and Permian macrofossil record, although there is palynological evidence of these plants having grown earlier in extra‐basinal habitats and was dominated by gymnosperms with more modern affinities. The Cenophytic Flora that first appeared during Cretaceous times was overwhelmingly dominated by angiosperms. The end‐Devonian, end‐Triassic and end‐Cretaceous mass‐extinction events recognized in the marine fossil record had little impact on the diversity dynamics of these Evolutionary Floras. Rather, the changes between floras mainly reflect key evolutionary innovations such as heterospory, ovules and angiospermy.     

Lower Devonian spore assemblages from the Arbuthnott Group at Canterland Den in the Midland Valley of Scotland

The Canterland Den locality in the Midland Valley of Scotland exposes sediments belonging to the Arbuthnott Group. They consist of typical Lower Old Red Sandstone terrestrial fluviatile deposits, which probably accumulated in an inland intermontane basin. Palynological investigation of these sediments has yielded diverse and well preserved palynomorph assemblages dominated by land-derived forms: spores, phytodebris (dispersed cuticles and tubular structures) and rare fragments of arthropod cuticle. Interestingly, rare acritarchs interpreted as deriving from non-marine algae are also present. The spore assemblages all belong to the lower subzone of the micrornatus–newportensis Spore Assemblage Biozone, indicating an early Lochkovian (Early Devonian) age. They are similar in general characteristics to previously described spore assemblages from the Arbuthnott Group, but exhibit minor differences in terms of taxon composition. They differ more significantly from coeval spore assemblages from the lowland floodplain deposits of the Anglo-Welsh basin, in terms of both relative abundance of morphotypes and taxon composition. An intriguing feature of the assemblage is the high abundance of undissociated spore tetrads. It is suggested that such tetrads are a genuine feature of Lochkovian spore assemblages, perhaps reflecting more flexible and unconventional reproductive strategies exhibited by early land plants.     

A diverse assemblage of early land plants from the Lower Devonian of the Welsh Borderland

FANNING, U., EDWARDS, D. & RICHARDSON, J. B., 1992. A diverse assemblage of early land plants from the Lower Devonian of the Welsh Borderland. Nine rhyniophytoid taxa are described from an early Gedinnian locality ( middle micromatus-newportensis spore Biozone) near Ludlow, England. They include Cooksonia pertoni, C. hemisphaerica, C. cambrensis, Tortilicaulis transwalliensis and three new taxa, Salopella marcensis sp. nov., Uskiella reticulata sp. nov. and Tarrantia salopensis gen. et sp. nov. Isolated sporangia of reniform shape, and those subtended by short lengths of axis, contain spores of Apiculiretusispora type and may belong to C. caledonica or Renalia . Morphologically distinctive forking, terminal sporangia lacking identifiable spores are not placed in a new taxon, because evidence based on in situ spores from elsewhere suggests they may belong to Salopella . Structures previously interpreted as clusters of sporangia of Yarravia- type are shown to be ± globular sporangia longitudinally split into valves. Sterile axes are dominated by smooth forms; although rare examples possess small enations. Tracheids have not been seen in axes of fertile specimens nor in sterile coalified compressions. A single pyrite permineralization contains tracheids of zosterophyll type. The assemblage demonstrates diversity among rhyniophytoids in the early Devonian and the existence of low vegetation 'alongside' the much larger zosterophyll dominated type.     

Tetralete and Trilete Spores in Adiantum Hispidulum (Adiantaceae) in New Zealand

Both tetralete and trilete spores have been found in New Zealand material of A. hispidulum Swartz. The rare tetralete spores are associated with the occurrence of spore pentads and hexads (post meiosis), and have the same sculptural pattern (finely pitted) as the trilete spores but are smaller with laesurae being often less distinct.     

Microfossil Assemblages from Upper Carboniferous in Ordos Basin,Northwest China

This paper is resulted from a systematic study of sporo-pollen of Ordos Basin of Northwest China has been found. Single spore assemblage in the Taiyuan Formation contains abundant Pteridophic and Pteridosperm spores and a few Gymnosperm pollen. 11 new species are described. New sporopollen taxa include, Punctatisporites gansuensis, Auritulinasporites ningxiaensis, Gulisporites discersus, Trivolites scabratus, Psomospora anulatus, Gravisporites rugu- laris, Velamisporites breviradidlis, Murospora strialatus, Foveomonoletes foveolatus, Cy- clogranisporites microtriagulus (comb. n.), Cycadopites pachyrrhachis (comb. n.) and Punatatisporites sp., Tantillus sp., Converrucosisporites sp., Columnisporites sp.. and Zono- spheridium reticulatoides and Z. granuloides of Acritarchs. There are some differences between spore assemblages of eastern and western parts of Ordos Basin. Western part is characteristic by more abundance of cingulum spores. The age of the microfossil is considered to be of the Late Carboniferous.