Physiological correlates of the morphology of early vascular plants

RAVEN, J. A., 1984. Physiological correlates of the morphology of early vascular plants. The early evolution of vascular land plants is considered in relation to the physiological problems of life on land. The universal characteristics of vascular plants (xylem, cuticle, stomata, intercellular air spaces, long-distance symplastic transport and alternation of generations) are discussed in terms of the essential properties of a homoiohydric phototroph. Likely precursors of vascular plants, and the physico-chemical and biotic environment in which they occurred, are outlined prior to a discussion of the selective forces acting on the evolution of vascular plants in the Upper Silurian and Lower Devonian. Emphasis is placed on biochemical and structural 'pre-adaptations' which may have occurred in the precursors of vascular plants and on which natural selection could have acted with lignified xylem, stomata, etc., as the end-products. Guiding principles in the analysis include the physiology of extant plants, physico-chemical constraints, and compatibility with the fossil record. It is concluded that the likely sequence of acquisition of vascular plant characteristics was: heteromorphic alternation of generations with an erect sporophyte; cuticularization of sporophyte; evolution of xylem; occurrence of intercellular air spaces with pores in the epidermis; stomatal activity of the pores. Endodermis and phloem-type long-distance transport probably originated around stages (3)-(5).     

Microplankton changes through the early Silurian Ireviken extinction event on Gotland, Sweden

This paper documents and analyses the extinction and origination patterns of acritarchs and prasinophyte algae at the Llandovery/Wenlock boundary transition in the Lower Silurian on the island of Gotland, Sweden. Closely spaced samples were collected from two parallel sections: Lusklint 1 and Lickershamn 2, spanning the upper part of the Lower Visby Beds and almost all of the Upper Visby Beds (i.e. the uppermost Llandovery and lowermost Wenlock).

At least eight extinctions affecting the conodont record have been reported at these levels. This turnover (named the Ireviken Event) has been interpreted as an example of the change from a P to an S climate state, reflecting large changes in the ocean/atmosphere system.

The palynomorph data show a significant turnover in the phytoplankton, with most of the extinctions at the end of the event (85.3% in the top 4 m of the Lusklint 1 section). The originations are more numerous than the extinctions and they are distributed through the whole of the Ireviken Event. There is an uneven distribution across the event with more originations in the Lower Visby Beds forming a convex pattern.

Comparison of these data to other palynological studies suggests that there was a slightly higher number of migrations out of Gotland than in. The P and S model does not successfully explain all the changes recorded, but is the model most inclusive of all the climatic variables available at this time.     

Genetically controlled cortical tissue deposition inNormalograptus scalaris (Hisinger, 1837)

A specimen ofNormalograptus scalaris (Hisinger) from the Llandovery of Dalarna, Sweden, shows a highly asymmetrical rhabdosome in which one stipe was abandoned at an early stage in the growth of the colony. The surface of the rhabdosome is thickly covered by cortical tissue, but the remaining dorsal side of the second stipe is left free. Even though the symmetry of the colony is strongly distorted, it survived for a considerable time. It is suggested that this unusual development involved genetic control of cortical tissue deposition in the graptolite colony.      

Preservation of soft tissues in Silurian graptolites from Latvia

The contractile stalks of graptoloid zooids are preserved as organic carbon residues in thecae of the middle Llandovery graptoloid graptolites Rastrites geinitzii and Neolagarograptus? sp. from the Aizpute‐41 core, Latvia. The contractile stalks are surrounded by equant pyrite crystals, resulting in three‐dimensional preservation of the graptolite rhabdosomes, and are associated with sediment of similar composition to, and derived from, the adjacent matrix. Matrix entered the thecae after pyrite crystal growth and filled some of the space left by collapse of the contractile stalks and some intercrystalline cavities; other space is partially infilled by diagenetic minerals. The contractile stalks are parallel‐sided and occupy up to one‐half the metathecal width, which is not inconsistent, assuming post‐mortem shrinkage, with the suggestion that graptoloid zooids filled their thecal tubes in life. The location of the preserved soft tissues, towards the distal ends of the metathecae, is very different from that predicted by decay experiments on the extant pterobranch hemichordate Rhabdopleura; the latter's soft tissues may thus not be a reliable analogue for those of these Silurian graptoloids.     

A sequence stratigraphical model for the Late Ludfordian (Silurian) of Gotland,Sweden: implications for timing between changes in sea level,palaeoecology, and the global carbon cycle

This paper investigates a time interval within the Late Ludfordian (Late Silurian), involving changes in faunal composition (the Lau Event), a major positive carbon isotope excursion (CIE), and contemporaneous sea-level changes in remote palaeo-basins. Based on the Silurian strata of Gotland (Sweden), we integrate sequence stratigraphy, carbon isotope stratigraphy, and platform-scale palaeoecological changes associated with this turbulent time period in Earth history. Three depositional sequences (sequences Nos. 1–3), including two separate periods of forced regression (falling stage systems tracts, FSSTs) are identified from outcrop and drillcore studies. The sequence stratigraphical framework is interpreted to reflect glacio-eustatic sea-level changes. The CIE starts at the onset of the initial FSST (sequence No. 1), just below the last appearance datum of the conodont Polygnathoides siluricus. The values increase through the ensuing lowstand and transgressive systems tracts (LST and TST) of sequence No. 2 and peak in the following highstand systems tract (HST). A second forced regression (FSST of sequence No. 2) took place in the lower Ozarkodina snajdri Zone. δ¹³C data are scarce from these siliciclastic strata, but inferably remain high. The δ¹³C values increase within the LST and earliest TST of sequence No. 3, before a decreasing trend starts within the early TST. δ¹³C values return to pre-excursion levels within the ensuing HST. The CIE is closely associated with an increase in stromatolites (mats and oncoids) across a wide range of depths and sedimentary environments, and correlations to other basins indicate a global increase in cyanobacterial activity. A drastic decline in level-bottom benthic faunas during the FSST of sequence No. 2 is, however, interpreted as a local response to the progradation of a delta complex (the Burgsvik Sandstone). Biological carbonate production replenishes rapidly within the TST of sequence No. 3, succeeding a thin LST dominated by reworked siliciclastics and chemically precipitated carbonates (ooids). The detailed relationship between the CIE and sea-level change presented herein is not fully consistent with previous reports on the CIEs associated with the lower Silurian Ireviken and Mulde events, respectively. Based on our facies analysis and sequence stratigraphical interpretation, two main mechanisms are suggested as responsible for the Late Ludfordian CIE: (1) a change in the riverine C-weathering flux towards the ¹³C end member following glacio-eustatically induced subaerial exposure of carbonate platforms throughout the tropics, and, (2) increased photosynthetic activity by benthic cyanobacteria exaggerating the δ¹³C values of precipitated carbonates.     

Peridermal and interthecal tissue in Silurian retiolitid graptolites: with examples from Sweden and Arctic Canada

Peridermal tissue, and thecal 'floors' are shown to be present in some specimens of Pseudoretiolites, Retiolites and Stomatograptus. The development of a dense reticulum and thecal hoods is also recognized as being a progressive developmental feature in Gothograptus. It is suggested that all these structures, whether internal or external, are the product of mature or old-age development. It appears questionable whether the Rhabdopleura model of rhabdosomal development adequately explains late-stage membranes and other adventitious tissues such as discussed herein.     

Late Ordovician palaeoceanographic changes as reflected in the Hirnantian–early Llandovery succession of Jämtland, Sweden

A study of the Upper Ordovician–Lower Silurian strata in Jämtland, central Sweden, shows that large-scale changes in shelf deposition took place close to the systems boundary. These changes include unconformity development and the replacement of a siliciclastic shelf with a carbonate-dominated shelf, suggesting the interaction of allocyclic controls such as changing eustatic sea-level and climate. The 6-m-thick Ede Formation is a key lithosome for interpretation of this transition. Its sediments were deposited in the Caledonian foreland basin, situated east of the closing Iapetus Ocean on the western margin of the Baltic craton. A major part of the late Caradoc to late Ashgill (into the Hirnantian) was characterised by continuous and uniform deposition over wide areas (Kogsta Formation), whereas erosional surfaces and complex lateral facies relationships characterise the Ordovician–Silurian boundary strata (Ede Formation and lateral equivalents). The Ede Formation represents the end of terrigenous deposition, which in the middle Aeronian was followed by regional expansion of carbonate deposition (Berge Formation). A syn-sedimentary erosional surface, with at least 1 m of relief locally, forms the lower boundary of the Ede Formation. This surface is overlain by two types of conglomerate. Lower parts of the Ede Formation consist of medium to thick-bedded quartzites. A second erosional surface with only minor (few centimetres) relief occurs on top of these quartzites. The upper parts of the Ede Formation consist of a thin, basal favositid biostrome overlain by thin bedded, calcareous sandstones, limestones and intensely bioturbated shales. Analysis of stratigraphic boundaries and the facies succession suggests that the lower Ede Formation represents a major downward shift in coastal onlap and by-pass sedimentation that created the lower erosional surface. The erosional surface in the middle of the Ede Formation is inferred to have formed during the subsequent maximum lowstand or as a ravinement surface, and is interpreted as an unconformity. The succession is subdivided into four facies associations, each corresponding to a specific systems tract: (a) a Shale–Siltstone Association (uppermost Kogsta Formation), deposited during a highstand situation in mid-outer shelf areas; (b) a Quartzite Association (the lower Ede Formation), deposited during forced regression in a shoreface environment; (c) a Mixed Carbonate–Siliciclastic Association (the upper Ede Formation), deposited during transgression in a wave-dominated, proximal shelf environment when clastic supply was reduced; and (d) a Micritic Limestone Association (lowermost Berge Formation), deposited during a second highstand situation in a low-energy, offshore environment.

Conodont data, together with a previously reported Hirnantia fauna, constrain the position of the Ordovician–Silurian boundary to the lower 1.65 m of the Ede Formation, or less likely, to the uppermost metre of the underlying Kogsta Formation, i.e., within a 2.65-m-thick uncertainty interval. The base of the Berge Formation is about 4 m above the top of the uncertainty interval, and is dated as being mid-Aeronian in age, suggesting condensation and/or a hiatus close to, or at, the Ordovician–Silurian boundary. These data tie the unconformity and the regional facies change from a siliciclastic to a carbonate-dominated shelf to Late Ordovician–Early Silurian eustatic and climatic changes.     

EARLY LLANDOVERY CHITINOZOANS FROM JORDAN

Abstract:  Chitinozoans recovered from cores BG-14 and WS-6 in southern and eastern Jordan are described. Sample ages are well-constrained by graptolite data to the ascensus-acuminatus and vesiculosus graptolite biozones. Eighteen chitinozoan taxa are recognized, with Spinachitina  fragilis , Plectochitina  nodifera , Ancyrochitina  laevaensis and Belonechitina  postrobusta allowing correlation with both local and global chitinozoan and graptolite biozonations for the Rhuddanian. One new species of chitinozoan is erected, Belonechitina  pseudarabiensis sp. nov.     

Microplankton associations, biofacies and palaeoenvironment of the type lower Ludlow Series, Silurian

The acritarchs and prasinophyte algae from the type lower Ludlow Series of the Goggin Road section, Ludlow, England, are resolved into seven recurrent associations comprising taxa with similar environmental preferences. Endemic and environmentally sensitive associations of acritarchs and prasinophytes are identified and high-resolution fluctuations in the early Ludlow palaeoenvironment are established. An early Ludlow crisis in the acritarchs is recognized in the lower part of the Middle Elton Formation, when an abrupt palaeoenvironmental change in the Ludlow area resulted in a large decline in the abundance of the acritarchs, but allowed Tasmanites and retiolitid graptolites to flourish briefly. Cymbosphaeridium sp. A, Pulvinosphaeridium ludlowense and Multiplicisphaeridium arbusculum forma A are taxa possibly specialized, or produced as a response, to a stressed palaeoenvironment, as they are most abundant when other acritarchs and prasinophytes are uncommon. The low abundance of acritarchs and prasinophytes in the Upper Elton Formation may be related to high sedimentation rates and to the slumping of sediments caused by instability on the shelf of the Welsh Basin, or to lower plankton productivity.