Tetrads in sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland

Prior to the mid-Silurian, evidence for the earliest embryophytes comes from dispersed spores, particularly permanent tetrads, there being no fossils showing gross morphology or anatomy of the producers. The fragmentary coalified mesofossils described here from the uppermost Silurian (Pridoli) and basal Devonian (Lochkovian) of the Welsh Borderland contain tetrads assigned to Tetrahedraletes, Velatitetras and Cheilotetras. These spores together with examples from spore masses have been examined by scanning and transmission electron microscopy and display diversity in ultrastructure of the exospore and envelope. Tetrads have been found, together with a putative elater, in the forking apex of an axial Lochkovian fossil, named Grisellatheca salopensis gen. et sp. nov., that anatomically, apart from spore characters, reveals no unequivocal evidence for hepatic affinity. The remaining fossils are equally as uninformative as regards affinity. Tetrads with ornamented envelopes are recorded in an isolated discoidal sporangium and in the bases of incomplete sporangia borne terminally on a bifurcating axis. Both ornament and ultrastructure suggest that the spores belong to quite distinct species within Velatitetras. Tetrahedraletes is recorded in an incomplete sporangium subtended by a forking axis, in which no cellular detail is preserved. Naked unfused tetrads also assigned to Tetrahedraletes are recorded in spore masses from both localities and again exospore ultrastructure demonstrates diversity. A final Lochkovian sporangium contains naked tetrads with sporadic Papiculate ornament and shows a unique trilayered exospore. Comparisons of exospore ultrastructure in these tetrads, which it is argued are mature and dispersed as such, provide no unequivocal evidence for affinities, be they tracheophyte or bryophyte. The bifurcating sporophytes are evidence against similarities with extant bryophytes. It is concluded that these very fragmentary fossils derive from small plants comprising relict populations of the vegetation that flourished on land in turfs through the greater part of the Ordovician and early Silurian, but that was gradually replaced by the tracheophytes.     

Permanent dyads in sporangia and spore masses from the Lower Devonian of the Welsh Borderland

Dyads, interpreted as cryptospore permanent dyads, are reported from sporangia and spore masses recovered from fluviatile deposits of Lochkovian (Early Devonian) age from the Welsh Borderland. The morphology, anatomy and ultrastructure of the specimens have been analysed using LM, SEM and TEM. Two specimens consist of a single sporangium terminating an axis: one is cup-shaped and attached to an unbranched axis and named Culullitheca richardsonii gen. et sp. nov., the other comprises an isotomously branching axis with one sporangium preserved and is named Fusitheca fanningiae gen. et sp. nov. Two specimens show dyads adhering to cuticular fragments presumably derived from sporangia. In one the cuticle is of irregular shape, in the other it is elliptical. The final specimen comprises an elongate spore mass. In all cases ultrastructural analysis reveals that the spore walls are essentially homogeneous. Our findings suggest that cryptospore permanent dyads were produced by rhyniophytoid plants of small stature. The affinities of these plants remain conjectural, as is the relationship of the cryptospore permanent dyads with other sporomorph morphotypes (cryptospores and trilete spores). However, the production of dyads by a plant with a bifurcating sporophytic axis is evidence against affinity with extant bryophytes. The mode of formation, adaptive significance, affinities, phylogenetic relationships and stratigraphical history (including demise) of cryptospore permanent dyads is discussed.     

A new genus for isolated bivalved sporangia with thickened margins from the Lower Devonian of the Welsh Borderland

A new taxon Sporathylacium salopense gen. et sp. nov . is based on small isolated coalified sporangia from the Lower Devonian (Lochkovian: micrornatus—newportensis spore zone) from the Welsh Borderland. The sporangia have two equal valves with multi-layered walls and thickened borders, and contain trilete crassitate, non-curvaturate isospores that are completely covered by a microgranular ornament with possible distal verrucate/murornate structures. They differ from zosterophyll sporangia in details of the presumed dehiscence zone extending around the entire free convex margin, particularly in the presence of a wedge of amorphous material between the valves, and in spore characters. Absence of any information relating to water-conducting cells prevents further assignment within the embryophytes. Spherical bodies associated with spores and a resilient sporangial lining are compared with similar structures in extant free-sporing plants and with Ubisch bodies. The mode of sporangial dehiscence involving anatomical modifications of the valve margins and the novel wedge of tissue which connects them remains speculative.     

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.     

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.     

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.     

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.     

Two cryptospore-bearing land plants from the Lower Devonian (Lochkovian) of the Welsh Borderland

Two examples of fragmentary, coalified plant fossils with cellular preservation and in situ spores are described using scanning and transmission electron microscopy (SEM and TEM), from a Lower Devonian (Lochkovian) locality in the Welsh Borderland. A sporophyte which produced dyads of ?Cymbohilates of. horridus is unique in that stomata are numerous on the supporting axis, and are comparable with those described from contemporary vascular plant remains. The in situ dyads possess a bilayered exospore wall, with an outer exosporal envelope present over the distal faces. A fusiform sporangium, with externally smooth epidermis, contains specimens of the tetrad Velatitetras sp. Each tetrad is encompassed by a laevigate, folded, exosporal envelope of uniform thickness, which contains a layer of regular voids. Spores within the tetrads are ultrastructurally bilayered, with a complex, digitate outer margin presumably representing spore wall ornamentation. Neither the tetrads nor dyads reported in this paper are lamellate in ultrastructure. The combination of stomata, branching and dyads in the same sporophyte holds significance for the understanding of cryptospore affinities.     

Fossil plants with spores in the sporangia from the Upper Devonian (Frasnian) deposits of northern Timan

Fossil plants from the Upper Devonian (Frasnian) of northern Timan are studied. The sporangia contain well-preserved spores, which were studied in transmitted light using a scanning electron microscope. The genus Gutzeitia S. Snigirevsky, gen. nov. is established. Macroremains and in situ spores of G. timanica (Petros.) S. Snig., comb. nov. and ?Cephalopteris mirabilis (Nath.) Nath. are described. Microspores in Dimeripteris gracilis Schmalh. are identified, and microspores and megaspores in ?Cephalopteris mirabilis are studied for the first time.     

An unusual microfungus in a fungal spore from the Lower Devonian Rhynie chert

Abstract: A new fossil microfungus, Kryphiomyces catenulatus gen. et sp. nov., occurs as an endobiotic mycelial thallus in a large spore of a glomeromycotan fungus from the Lower Devonian Rhynie chert. The thallus consists of branched (?pseudo‐)septate hyphae with numerous catenulate swellings. Some hyphal tips produce spherical reproductive structures or propagules. Hyphal morphology in K. catenulatus is reminiscent of that in certain extant Hyphochytridiomycota, Chytridiomycota, and even Ascomycota, but specific diagnostic features that allow assignment of the fossil to modern groups are absent. The discovery of this interfungal association broadens our knowledge about the diversity of microfungi and their intricate associations in early continental ecosystems.     

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.     

Upper Ordovician and Lower Silurian chitinozoans from central Nevada and Arctic Canada

Late Ordovician and Early Silurian chitinozoans from the uppermost Vinini Formation, and the Hanson Creek Formation in central Nevada and the lower Cape Phillips Formation, Cape Manning section, in Arctic Canada have been re-investigated and a new chitinozoan biozonation is proposed. The Upper Ordovician of central Nevada can easily be correlated to that of Arctic Canada through the common occurrence of the Ordochitina nevadensis biozone in both regions. No such correlation, however, is possible with the Late Ordovician of Anticosti Island in eastern Canada because of the absence of the index Upper Ordovician chitinozoan zonal species of central Nevada such as Belonechitina tenuispinata sp. nov, Ordochitina nevadensis sp. nov. and Nevadachitina vininica gen. nov., sp. nov. in the former area.One new genus, Nevadachitina, and nine new species, Eisenackitina ripae, Belonechitina martinica, Nevadachitina vininica, Nevadachitina praevininica, Ordochitina nevadensis, Belonechitina tenuispinata, Belonechitina parvispinata, Tanuchitina laurentiana, Angochitina hansonica are described and illustrated in this paper and four species are left in open nomenclature.     

Caradoc strophomenoid and plectambonitoid brachiopods from Wales and the Welsh Borderland

Abstract: Ordovician strophomenoidean and plectambonitoidean brachiopods are reviewed and partially redescribed from the Caradoc (Sandbian to Early Katian) age rocks of Wales and the Welsh Borderland of England, then forming part of the Avalonia Terrane. There are 51 nominal species available, of which 12 are synonymised here and many transferred to different genera. Strophomenoids occurred sporadically, often rarely but occasionally abundantly, particularly in mid‐shelf benthic assemblages, and 21 named species and subspecies are noted, including the new Kiaeromena (Kiaeromena) harperi, as well as six in open nomenclature. The common genus Kjaerina, with six species known from the area, is revised in detail. In contrast to the strophomenoids, plectambonitoids were more common in shallow and mid‐shelf assemblages, including Eoplectodonta (Eoplectodonta) abigailae sp. nov., with Sowerbyella in particular dominating many bedding planes, and 17 named species are reviewed, as well as three in open nomenclature. The distinctive Gunningblandella, hitherto known only from Australia and Kazakhstan, is recorded for the first time from Europe. The Avalonian strophomenides are compared with those from neighbouring terranes, particularly Laurentia and Baltica, during a time when the seas between those terranes were narrowing, and the faunas from the three terranes are found to be largely similar at the generic level, although there are few species in common. In contrast, there are few genera in common between those three terranes and the Mediterranean Province of Northwestern Gondwana.     

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.     

Induced release of Bacillus spores from sporangia by sodium sulphate

Incubation of sporulating cultures of Bacillus anthracis, B. cereus, B. subtilis and B. thuringiensis in 1.0 mol/l sodium sulphate markedly increased the release of free spores from sporangia. It is postulated that the release of spores is due to activation of latent autolysins which hydrolyse sporangial cell walls. Sodium sulphate-induced lysis of sporangia represents a novel and highly effective method for the recovery of spores from cultures of Bacillus species.     

Wenlock carbonate,level bottom,brachiopod-dominated communities from Wales and the Welsh Borderland

Four marine level bottom communities are described from Wenlock carbonate rocks (i.e. the Wenlock and Woolhope limestones) of Wales and the Welsh Borderland. The communities are: (1) Sphaerirhynchia wilsoni, (2) Isorthis clivosa, (3) Eoplectodonta duvalii, and (4) Visbyella trewerna. The Sphaerirhynchia wilsoni community is divided into: (a) an argillaceous micrite phase, and (b) a biomicrite phase. The communities and the phases are defined on the basis of consistent and recurrent numerical species combinations.The community spectrum from Sphaerirhynchia wilsoni to Visbyella trewerna is broadly correlated with an increase in water depth. The communities show no correlation with major limestone facies types. It is suggested that turbulence, rates of sedimentation and substrate type are some of the environmental parameters, in the shallow near-shore regime, which determine the distribution of brachiopods.     

New bryozoans (Bryozoa) from the Upper Devonian (Lower Famennian) of the Kuznetsk Basin

New bryozoans from the Upper Devonian (Famennian) of the borderlands of the Kuznetsk Coal Basin are described: Orthopora tomensis sp. nov. and Minussina incrustata sp. nov.