Miospore assemblages from the Silurian–Devonian boundary,in borehole A1-61, Ghadamis Basin,Libya

Well preserved and diversified miospore assemblages have been recorded from a relatively continuous sequence in borehole A1-61 which spans the Silurian–Devonian boundary in the northwestern part of the Ghadamis Basin, Libya. The sequence is represented by early Devonian Lochkovian beds of the Tadrart Formation that transgress onto the Silurian Ludlow-Pridoli beds of the upper part of the ‘Alternances Argilo-gréseuses’ Formation. The present work demonstrates a succession of miospore assemblages from closely sampled layers that have been stratigraphically dated as Ludlow–middle Pridoli and early Lochkovian by chitinozoans and acritarchs. Over 80 species of cryptospores and trilete spores have been identified. Modified detailed morphological interpretations are given. The miospore assemblages are correlated with miospore zonation schemes established for the type sequences of the Welsh Borderland, and those previously described from Libya. Early occurrences of some species as Streelispora newportensis on the western Gondwana plate, are put forward by comparison with the Old Red Sandstone continent. Phytogeographic and palaeobotanic implications based on these observations are discussed.     

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。     

A new plant assemblage (microfossil and megafossil) from the Lower Old Red Sandstone of the Anglo-Welsh Basin: its implications for the palaeoecology of early terrestrial ecosystems

Lower Old Red Sandstone deposits penetrated by a series of cored boreholes near Newport (South Wales) have been sedimentologically logged, and recovered plant assemblages (microfossil and megafossil) investigated. Sedimentological logging indicates that the deposits are typical of the extensive terrestrial-fluviatile floodplain deposits of the Anglo-Welsh Basin. Palynomorph assemblages have been recovered from a number of horizons and comprise entirely terrestrial forms (spores and phytodebris). They essentially represent a single assemblage, belonging to the middle subzone of the micrornatus-newportensis sporomorph assemblage biozone, and indicate an Early Devonian (mid-Lochkovian) age. The new biostratigraphical data enables correlation with other Lower Old Red Sandstone deposits of the Anglo-Welsh Basin, and the deposits are assigned to the lower part of the St. Maughan's Group. A plant megafossil/mesofossil assemblage recovered from one of the spore-bearing horizons includes a zosterophyll assigned to Zosterophyllum cf. fertile. This is the earliest reported zosterophyll from the Anglo-Welsh Basin. The new palynological/palaeobotanical data provide important information on the palaeoecology and palaeobiogeography of the vegetation of the southeastern margin of the Old Red Sandstone continent during Lochkovian times. Palaeogeographical variation in the distribution of plant microfossils and megafossils is interpreted as reflecting differences between the flora of the lowland floodplain and inland intermontaine basins, although this is to a certain extent overprinted by variation due to localized differences in environmental conditions.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Siluro-Devonian miospore biostratigraphy of the Urubu River area, western Amazon Basin, northern Brazil

Well-preserved miospores have been observed in the Urubu River area from the western part of the Amazon Basin, northern Brazil, in rocks belonging to the uppermost Pitinga and Manacapuru Formations. A previous palynological work based on chitinozoans has dated these strata as Late Ludlow, Early Pridoli and Early Lochkovian. Although samples are relatively poor in miospores, the present study reveals the most diversified Silurian-Earliest Lochkovian palynoflora ever reported from South America, an interval where miospore information is still very rare. Sixty-four species have been identified, of which two are new (Artemopyra urubuense and Retusotriletes amazonensis). One single specimen attributed to the genus Grandispora, identical to unpublished forms identified by one of us (P.S.) in Lochkovian sections of Saudi Arabia, has been found in a Lochkovian sample from the Urubu River. These constitute the oldest geological record of Grandispora. Among the abundant acritarchs, specimens of the genus Schizocystia have been observed for the first time in pre-Devonian rocks.Although the Late Silurian-Early Lochkovian miospore biostratigraphy is still in its infancy in South America, it seems that Amazon Basin assemblages do not show significant differences in composition in comparison to coeval palynofloras from other areas of northwestern Gondwana. This is surprising since the miospore localities under consideration are widely distributed over high to low paleolatitudes, i.e. from cool temperate to dry subtropical belts. Conversely, Siluro-Devonian Gondwanan assemblages differ considerably from the contemporary palynofloras of the Old Red Sandstone Continent and adjacent areas, thus contradicting recent phytogeographic interpretations based on megafloras.     

Miospore evolution from the Ordovician to the Silurian

The definition of the term cryptospore is amended to include only spores thought to be produced by embryophytes and to exclude all enigmatic palynomorphs. Cryptospores are included here in the miospore group.The oldest published assemblage of typical cryptospores is Llanvirn in age. The evolution of cryptospore assemblages is very slow from the Llanvirn to the early Llandovery. The Telychian and Sheinwoodian stages are periods of drastic impoverishment in cryptospore biodiversity, followed by an important modification in miospore assemblages. This step in the evolution of the vegetation is correlated with the rapid transgression of seas on the continent during the early Llandovery. On the other hand, the Hirnantian glaciation does not affect diversity of the miospore assemblages.     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age ( micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.     

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.     

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.     

Aspects of Gondwana paleobotany: gymnosperms of the Paleozoic—Mesozoic transition

During the late Paleozoic and early Mesozoic the Gondwana Supercontinent underwent dramatic geographic and climatic changes. Geologic and biologic factors concurrently played an important role modelling the vegetation of that time. The gymnospermic component of plant assemblages shows significant variations in composition and in the distribution of different taxa. Analysis of the assemblages shows that some plant groups dominated the scenario, such as the pteridosperms, glossopterids, corystosperms and, to a lesser degree, cordaites and conifers. Ginkgophytes, bennettites or cycads were less important in the Paleozoic but their numbers increased in the Triassic. Paleozoic assemblages were extensively dominated by glossopterids that became extinct in the earliest Mesozoic. Pteridosperms crossed the P—M barrier and became dominant during the Triassic, at a time when corystosperms evolved and radiated. Other groups became extinct in the Paleozoic, namely dicranophylls and cordaites. Conifers were represented by different families, restricted either to the Paleozoic or the Mesozoic. They were not conspicuous in the analysed assemblages. In some areas of Gondwana, taxa of the Euramerican alliance are present through a migrational mechanism that occurred during continental displacements which produced global climatic changes. Recent studies have shown that there are far more common elements between Euramerica and Gondwana than suspected up to now. These elements find their distribution especially in the western part of Gondwana (Africa—South America).     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age (micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.     

Environmental changes reflected by palynomorphs in the early Middle Ordovician Hanadir Member of the Qasim Formation, Saudi Arabia

Core samples from the QASIM-801 (QSIM-801) water well in the Qasim Region of central Saudi Arabia were investigated palynologically. The interval studied contains the transgressive succession from the Sajir Member of the Saq Formation to the Hanadir Member of the Qasim Formation, and yields well-preserved and abundant cryptospores, microphytoplankton (acritarchs and chlorophycean algae), and chitinozoans, indicative of a Darriwilian (Llanvirn) age. The lowermost portion of the cored interval from the upper Sajir Member consists of fine-grained sandstones deposited in tidal flat, and shallow-marine settings. It is characterized by abundant and diverse cryptospores (permanent tetrads, dyads, monads), cuticle-like phytoclasts, and a low diversity assemblage of marine palynomorphs. The upper part of the cored interval from the Hanadir Member is composed of hemipelagic to pelagic, organic-rich, graptolitic shales, and micaceous shales, of middle- to outer-shelf depositional settings, which yield abundant marine palynomorphs (acritarchs and chitinozoans). Cryptospore-bearing horizons are interspersed in the succession. Changes in the composition of palynomorph assemblages reflect higher frequency environmental changes within an overall transgressive succession. Four new acritarch species are described in open nomenclature, Tyrannus sp. A, Clypeolus sp. A,?Pulvinosphaeridium sp. A, and?Tinacula sp. A.     

Pollen and spore assemblages in the Luwumbu Coal Formation (Lower Karroo) of the North Luangwa Valley,Zambia, and their biostratigraphic significance

Pollen and spore assemblages from the Luwumbu Coal Formation (Lower Karroo) of the northern part of the Luangwa Valley, Zambia, are described. Two assemblages have been recognised, one from banded siltstones and mudstones in the lower part of the formation, and the other from the overlying carbonaceous sediments of the upper part of the formation. The older assemblage is dominated quantitatively by monosaccate forms (Cannanoropollis and Plicatipollenites), and is generally similar to assemblages recorded by other workers from the Lower Karroo glacial and peri-glacial rocks of Zaire.The younger assemblages comprises a variety of supra-generic groups; the most common forms are disaccate (striate and non-striate) and trilete, but monosaccate, polyplicate, colpate, monolete and alete forms all occur. Genera which are especially common include Protohaploxypinus, Vesicaspora and Acanthotriletes. Cannanoropollis is abundant in the lower part of the younger assemblage. In general, the assemblage is similar to that recorded from the upper part of the Lower Coal Measures of the Ketewaka/Mchuchuma coalfield of Tanzania.