Biostratigraphy, biodiversity and palaeoenvironments of the chitinozoans and associated palynomorphs from the Upper Ordovician of the Central Anti-Atlas, Morocco

The range, abundance and diversity of Upper Ordovician chitinozoans and of various other organic-walled microfossils (i.e., acritarchs, leiospheres, scolecodonts) have been documented in the Bou-Ingarf section, Central Anti-Atlas, Morocco. The goal of this study is to provide a refined biostratigraphic scheme for the Lower Ktaoua, the Upper Tiouririne, the Upper Ktaoua and the Lower Second Bani formations in order to locate the initial effects of the Late Ordovician glaciation on the composition of assemblages of organic-walled microfossils. Five Late Ordovician chitinozoan biozones belonging to the North Gondwana zonation are identified. During the late Caradoc and early Ashgill, no significant changes in chitinozoan assemblages are recorded that could be attributed to any dramatic climatic and/or environmental events. The almost absence of phytoplanktonic elements (acritarchs and leiospheres) and of cryptospores in the fistulosa/barbata and in the nigerica biozones are interpreted as resulting of lower offshore environmental conditions. The reappearance of a variety of palynomorphs in the lower part of the elongata biozone does not indicate drastic climatic changes but more likely a shallowing trend of the marine environment and an increased primary production, possibly related to a cooling of the surface water masses subsequently to early effects of a general climatic cooling. The first obvious evidence of the Late Ordovician glaciation occurs in the lower part of the elongata biozone, at the base of the sandy member topping the Lower Second Bani Formation. This sandy deposit is interpreted as the result of a dramatic fall of the sea level due to the development of the Hirnantian ice cap on Gondwana. This short-lived glaciation ended when the northern Gondwanan ice cap melted during the upper part of the elongata and the oulebsiri biozones. A biological crisis affected both the chitinozoans and the acritarchs just after this glaciation, i.e., in the latest Hirnantian.     

Palynostratigraphy and palaeogeography of the Padeha, Khoshyeilagh, and Mobarak formations in the eastern Alborz Range (Kopet-Dagh region), northeastern Iran

A total of 113 surface samples collected from the Padeha, Khoshyeilagh, and Mobarak formations of Kuh-e-Ozum, northeast of Jajarm town were processed for palynomorphs, in order to determine age relationships. Well-preserved and abundant palynomorphs dominated by organic-walled-marine microphytoplankton (acritarchs and prasinophyte phycomata), miospores and subordinate chitinozoans, and scolecodonts were recovered. Seven species of prasinophyte phycomata (four genera), 19 acritarch species (14 genera), one species of chitinozoa, and 26 miospore species (19 genera) were recorded and assigned to eight local Assemblage Zones. Assemblage Zones I-IV occur in the Padeha Formation and suggest an Early Late Devonian (Frasnian) age whilst assemblages zones V-VII are present in the Khoshyeilagh Formation and indicate Late Devonian (Famennian) ages. Assemblage zone VIII, which occurs in the basal part of Mobarak Formation, suggests a Lower Mississippian (Tournaisian) age for this formation. Many of the palynomorph groups encountered are closely comparable with coeval assemblages recorded from Western Australia, southwest Ireland, England, Turkey, Saudi Arabia, North Africa, and South America, indicating the close relationship of the Iranian Platform to other parts of the northern Gondwana Domain during the time interval represented by these strata. The presence of marine palynomorphs (acritarchs/prasinophyte phycomata, chitinozoans, and scolecodonts), and shelly macrofauna (brachiopods, gastropods, and corals) in Member c of the Padeha Formation (as well as the Khoshyeilagh and Mobarak formations), together with associated miospores, indicate an open marine (moderately nearshore) depositional environment for the Upper Devonian and Lower Carboniferous deposits in northeastern Alborz Range (Kopet-Dagh region) of Iran.     

Upper Ordovician microphytoplankton of the Bill's Creek Shale and Stonington Formation,Upper Peninsula of Michigan,U.S.A.: Biostratigraphy and paleogeographic significance

An abundant, diverse, and well-preserved organic-walled microphytoplankton assemblage is described from the Upper Ordovician Bill's Creek Shale and the lower Stonington Formation (Bay de Noc Member) in the Upper Peninsula of Michigan, U.S.A. Based on graptolite and conodont evidence, the Bill's Creek Shale and Stonington Formation are Richmondian (=Ashgill) in age. The assemblage is dominated by acritarchs, which comprise 29 species (including the enigmatic palynomorph Gloeocapsomorpha prisca) assigned to 20 genera. The prasinophyte phycomata are represented by undifferentiated species of Leiosphaeridia and Tasmanites. In addition, chitinozoans are abundant, and scolecodonts and graptolite fragments are common. Paleontologic-palynologic and sedimentologic evidence indicates that the Bill's Creek Shale was deposited in a low-energy, shallow, nearshore marine environment. The overlying Bay de Noc Member of the Stonington Formation also accumulated in a low-energy, normal marine environment, but in a more offshore, somewhat deeper water setting. Both formations experienced minor transgressive and regressive episodes as indicated by fluctuations in the composition of the palynoflora. The combined Bill's Creek/Stonington acritarch assemblage closely resembles those described from the Richmondian-aged Maquoketa Shale (Missouri and Kansas), Sylvan Shale (Oklahoma), and Vauréal Formation (Anticosti Island, Québec, Canada). The overall composition of the acritarch assemblage from these four formations reflects a distinctive, recognizably Laurentian character. Nonetheless, many of the Bill's Creek/Stonington acritarchs have been reported from Upper Ordovician localities elsewhere, providing additional evidence for Late Ordovician cosmopolitanism of the marine microphytoplankton community. Additionally, the restricted stratigraphic range of many of the taxa further enhances their biostratigraphic application, both regionally and globally, and reaffirms the Richmondian (=Ashgill) age of the Bill's Creek Shale and Stonington Formation.     

The effects of the final stages of the Late Ordovician glaciation on marine palynomorphs (chitinozoans, acritarchs, leiospheres) in well Nl-2 (NE Algerian Sahara)

Palynomorph assemblages, especially chitinozoans and acritarchs, from the Upper Ordovician of well Nl-2 (north-east of the Algerian Sahara) are studied in order to precisely date the ultimate effects of the Late Ordovician glaciation and to document the impact of this major climatic stress on the diversity of the palynoplankton. References are made to stable isotope excursions and to global eustatic sea level variations in order to improve the local age determination. The Hassi el Hadjar Formation, i.e. glacio-marine diamictites, is interpreted as a transgressive event resulting from the melting of the northern Gondwana ice cap. It yields poorly preserved and moderately diverse chitinozoans of late Hirnantian age. Acritarchs are more abundant in the lower part of these "microconglomeratic clays", but display a low diversity and are badly preserved throughout the whole formation. Reworked individuals are recorded in both groups. The marine sediments of the M'Kratta Formation of latest Hirnantian age contain better preserved, more abundant and more diverse palynomorph assemblages, especially in the Upper Member. The composition of this palynoplankton indicates a fairly good faunal and phytoplankton recovery after the early Hirnantian climatic stress.The extinction of the Ordovician forms, and the appearance of Silurian type taxa occur only in the uppermost Hirnantian, i.e. following with a slight delay the glacial event. The overlying black shales of Wenlock age (lower part of the Oued Mehaiguène Formation) are indicative of marine anoxic environments. They yield a virtually exclusive, but enormous number of Tasmanacea. The very peculiar composition of this palynoplankton seems to be independent of the Late Ordovician glaciation and is most likely related to the factors that, later, generated and maintained anoxic conditions in this area.A new species of chitinozoan, Spinachitina oulebsiri sp. nov. from the latest Hirnantian M'Kratta Formation, is described and illustrated.     

Biostratigraphy,biodiversity and palaeogeography of late Silurian chitinozoans from A1-61 borehole (north–western Libya)

High resolution biostratigraphic investigation of well preserved and highly diversified chitinozoan assemblages from core samples collected in the upper part of the Alternances Gréso-argileuses Formation (late Silurian) in well A1-61 (Tripolitania, north–western Libya) is undertaken. The chitinozoan assemblages document the range of several Pridolian taxa selected as index species for the global Silurian chitinozoan biozonation. Distribution of important accompanying species is also discussed. The abundance and the diversity of these chitinozoans are quantified and collated with the local environmental trends as deduced from the lithology, sedimentary features and from the associated organic walled microfossils (i.e. scolecodonts, eurypterid fragments, acritarchs, spores, leiospheres, cuticles and plant debris). The palaeogeographic distribution of late Silurian chitinozoans is discussed with a particular attention paid to a very distinctive chitinozoan assemblage characterising the middle part of the Pridoli of the Ibarmaghian Domain in northern Gondwana regions. Systematic notes, including detailed biometric investigations and SEM illustrations, are provided for selected taxa of taxonomic interest and useful for regional or more long distance correlation.     

Palynostratigraphy of Middle Cambrian to lowermost Ordovician stratal sequences in the High Zagros Mountains, southern Iran: Regional stratigraphic implications, and palaeobiogeographic significance

A palynological investigation of Cambro-Ordovician stratal sequences in the High Zagros Mountains of southern Iran permits the definition of a series of successive acritarch assemblage zones of chronostratigraphic significance, much improving the current knowledge of the Lower Palaeozoic stratigraphy of this important area for oil exploration. The five acritarch assemblage zones can be readily correlated with previously established palynostratigraphic schemes constrained by co-occurrence of independent age evidence, confirming the utility of organic-walled microfossils for the detailed biostratigraphic characterization of sedimentary units. The proposed biozonation will facilitate accurate dating of the southern Iranian Cambrian sequences during future drilling of deep test oil wells. Acritarch assemblage zone I (Middle Cambrian), occurs at the base of Member C of the Mila Formation; assemblages zone II (late Middle to earliest Late Cambrian) extends through the middle and upper part of the same lithostratigraphic unit; zone III (early Late Cambrian in age) characterizes the lower part of the Ilebeyk Formation; zone IV (middle Late Cambrian up to Cambrian/Ordovician transitional levels) occurs in the middle and upper part of the Ilebeyk Formation; finally, acritarch assemblage zone V ranges through the basal part of the Zardkuh Formation and proves an early Tremadocian age for the latter unit. The Mid-Late Cambrian acritarch associations show a marked Avalonian palaeobiogeographical affinity, also sharing a high proportion of taxa with typical Baltican and North Africa–Gondwanan assemblages; on the other hand, they are clearly different from known Laurentian (North America) fossil microphytoplankton suites. These results are in general agreement with current palaeogeographical models which place Avalonia, Baltica, and the North African part of Gondwana, all at relatively high southern palaeolatitudes, in contrast with the sub-equatorial position of Laurentia. However, the presence of many typical “Avalonian” taxa in the Iranian Mid-Late Cambrian assemblages would suggest a closer position of Iran to Avalonia than currently envisaged. The observed breakdown of acritarch biogeographic differentiation in earliest Ordovician times possibly represents a major disruption of oceanic current patterns and a lessened palaeolatitudinal thermal gradient.     

Ordovician chitinozoan biozonation of the Brabant Massif, Belgium

Chitinozoans from seven Ordovician units (Abbaye de Villers, Tribotte, Rigenée, Ittre, Bornival, and Brutia formations and a new unnamed unit, here provisionally called the Asquempont unit) belonging to the mainly concealed Brabant Massif, Belgium are described herein. Fifty-six samples were taken from rocks cropping out at the south-eastern rim of the massif in the Orneau, Dyle-Thyle and Senne-Sennette valleys. Microfossil preservation is moderate to poor, and the chitinozoans occur in low numbers. Taxonomically, the recovered chitinozoans are distributed into 29 taxa, some placed under open nomenclature. Together with earlier published graptolite and acritarch data, the analysis of the chitinozoan assemblages resulted in an improved chronostratigraphy of the investigated formations. We propose a local chitinozoan biozonation with 11 zones for the Brabant Massif. The oldest investigated units yielded chitinozoans typical for North Gondwana, and younger units (starting in the middle Caradoc), yielded some taxa also common in Baltica. As the Brabant Massif formed part of the microcontinent Avalonia, the chitinozoan assemblages recovered from the massif support the inferred drifting of Avalonia from high latitudes towards middle latitudes in the Ordovician as was suggested earlier.     

塔里木盆地晚奥陶世良里塔格型生烃母质生物

本文描述塔里木盆地晚奥陶世塔中北斜坡带良里塔格组的生烃生物组合特征。指出缓斜坡背景下与灰泥丘有关的良里塔格组的生烃母质生物先质以混合型母质为特征。其组成主要有底栖宏观藻类、线叶植物、隐孢子、疑源类和几丁虫等。这是一类区别于世界上其它奥陶纪烃源岩Guttenburgen型的一类与灰泥丘有关的烃源岩的生烃母质生物类型。     

Infrared video microscopy—an efficient-method for the routine investigation of opaque organic-walled microfossils

Summary Dark or even opaque organic-walled microfossils are frequently represented in palynological residues which are derived from sediments of high maturity. Such particles may be studied surficially by SEM or incident light, but internal details remain obscured. It has been known for some time that transparency may be achieved by applying infrared microscopy, but the method never became routine. We have now assembled an IR equipment which is based on a regular transmitted light microscope and tested our setup on various kinds of palyno-morphse.g. spores, acritarchs, chitinozoans, scolecodonts as well as phytoclasts and zooclasts. Our examples are from several localities of different stratigraphic and palaeogeographic position within the Palaeozoic. The studies revealed a high potential for IR video microscopy to become an important tool in the study of thermally altered organic particles in palynological slides.     

The Ordovician Biodiversification: revolution in the oceanic trophic chain

The Early Palaeozoic phytoplankton (acritarch) radiation paralleled a long-term increase in sea level between the Early Cambrian and the Late Ordovician. In the Late Cambrian, after the SPICE δ¹³C_carb excursion, acritarchs underwent a major change in morphological disparity and their taxonomical diversity increased to reach highest values during the Middle Ordovician (Darriwilian). This highest phytoplankton diversity of the Palaeozoic was possibly the result of palaeogeography (greatest continental dispersal) and major orogenic and volcanic activity, which provided maximum ecospace and large amounts of nutrients. With its warm climate and high atmospheric CO₂ levels, the Ordovician was similar to the Cretaceous: a period when phytoplankton diversity was at its maximum during the Mesozoic. With increased phytoplankton availability in the Late Cambrian and Ordovician a radiation of zooplanktonic organisms took place at the same time as a major diversification of suspension feeders. In addition, planktotrophy originated in invertebrate larvae during the Late Cambrian–Early Ordovician. These important changes in the trophic chain can be considered as a major palaeoecological revolution (part of the rise of the Palaeozoic Evolutionary Fauna of Sepkoski). There is now sufficient evidence that this trophic chain revolution was related to the diversification of the phytoplankton, of which the organic-walled fraction is partly preserved.     

Acritarchs from the Dadas Formationin Southeast Turkey

Dadas Formation in Southeast Turkey yields abundant well preserved acritarchs, chitinozoans, tasmanites and scolecodonts. Encountered acritarchs suggest Upper Llandoverian-Ludlovian age for this formation. The acritarch association of the Dadas Formation resembles to the Middle Silurian assemblages of Tunisia and Libya. Thus Southeast Turkey falls in the Transitional-Iberian realm of Neoveryhachium carminae facies.     

Silurian calcispheres (Calcitarcha) of Gotland (Sweden): Comparisons with calcareous dinoflagellates

Scanning electron microscope examinations of polished and etched surfaces of sediments from the Silurian carbonate platform of Gotland, Sweden, revealed the presence of numerous, morphologically diverse “calcispheres” (Calcitarcha). Some of these spherical calcareous microfossils display wall structures that are surprisingly similar to those of calcareous dinoflagellate cysts. In analogy to the interpretation of the biological affinities of Palaeozoic acritarchs as cysts of organisms that might have been the ancestors of organic-walled dinoflagellates, the Calcitarcha from Gotland can be compared and may possibly be related to organisms that may have been the ancestors of calcareous cyst-producing dinoflagellates that so far have not been observed before the Late Jurassic.     

The environmental response of Middle Ordovician large organic walled microfossils from the Goldwyer and Nita Formations, Canning Basin, Western Australia

Middle Ordovician large organic walled microfossils (chitinozoans, scolecodonts, hydrozoans and foraminiferal linings) were recovered from the upper Goldwyer and lower Nita formations, Canning Basin, Western Australia, from three cores (WMC Santalum 1A, Kunzea 1 and Acacia 2). Petrophysical logs of these cores reveal an overall upward shallowing supersequence, overprinted by numerous transgression/regression couplets that can be correlated over 100km.Analysis of the abundance of the microfossils with respect to the gamma log signatures reveals that both chitinozoan abundance and diversity decrease as water depth shallows; however, the opposite is not always true and other factors probably intervene. Scolecodonts show an increase in abundance in transgressions, while hydrozoans and foraminiferal linings show no consistent response to trangressive or regressive phases. Cyathochitina hunderumensis tends to dominate chitinozoan assemblages where there is a transgression, while species of Belonechitina replace Cy. hunderumensis in regressive phases.     

Biodiversity patterns of Early–Middle Ordovician marine microphytoplankton in South China

Based on new materials from six sections and all available literature data, new diversity curves are presented for the phytoplankton (acritarchs) from South China, covering the Early–Middle Ordovician interval, when the Great Ordovician Biodiversification Event took place. The total diversity curve and the origination data imply that a major radiation of the phytoplankton occurred during the analysed interval. A peak of the total acritarch diversity curve appears in the A. suecicus graptolite biozone. The diversity changes vary in the different parts of the investigated area, most probably depending on the position of the analysed sections on the carbonate shelf or the slope, reflecting diversity differences due to the position on an inshore–offshore transect.The Early–Middle Ordovician diversity pattern of the phytoplankton is compared with those of several marine invertebrate groups. Compared with the diversity curve peak of the acritarchs, the conodonts and brachiopods reached their highest diversities before the acritarchs, while the highest diversity of the chitinozoans appears slightly later. The graptolites show two peaks during the Early–Middle Ordovician, while the trilobites diversity curve shows a peak only in the Sandbian. The different fossil groups, such as chitinozoans, conodonts, graptolites, brachiopods and trilobites show therefore different evolutionary patterns to that of the acritarchs, that are not yet fully understood, and correlations are so far difficult.The acritarch diversity changes can partly be compared to the local sea-level changes from four sections in South China. At a larger scale, the acritarch radiation coincides with a general transgression. At a regional or local scale, correlations are not straightforward, pointing out that more detailed data, based on both acritarch studies and more precise sea-level investigations, are necessary.     

新疆哈密地区晚泥盆世疑源类化石的发现及地质意义

从新疆哈密地区石城子北剖面7件样品获得分异度较高、有机质壁显著炭化保存的疑源类化石,根据其形态特征,共鉴定出14个形态属和15个形态种(其中8个未定种,3个比较种)。结合国内外古生代晚泥盆世已知疑源类组合进行比较,清楚表明当前获得的疑源类组合代表了晚泥盆世海洋微体浮游植物群面貌。我国涉及晚古生代疑源类生物地层的调查研究相对薄弱,而有关晚泥盆世疑源类化石的发现和报道更为匮乏,当前疑源类化石的发现,填补和丰富了新疆乃至国内晚泥盆世疑源类化石研究资料。该发现佐证了关于卡拉麦里洋在早石炭世闭合的认识;作为基础食物链的海洋微体浮游植物是重要成烃生物,晚泥盆世疑源类的保存预示研究区域具有石油、天然气勘察的前景。     

A SERIES OF MONOGRAPHS AND MEMOIRS (ISSN 0300-9491) TO SUPPLEMENT LETHAIA

Acid-resistant organic microfossils are described from outcrops and a boring through the middle and upper units of the VisingsÖ Beds in southern Sweden. 21 previously described acritarch species have been identified, and five new species are erected. The acritarchs include simple, although characteristically ornamented sphaeromorphs, one polygonomorph species and one acanthomorph species. Their mode of occurrence and possible lithofacial relationships are discussed. The acritarchs from the middle unit of the Visingso Beds indicate a Late Riphean age. The upper unit contains acritarchs previously known from Vendian and Lower Cambrian strata in the U.S.S.R., China and elsewhere. A Vendian age is suggested for this unit. A glacial origin is suggested for boulder beds in the middle unit of the VisingsÖ Beds.     

Ordovician and Silurian acritarch assemblages from the west Leinster and Slievenamon areas of southeast Ireland

The Lower Palaeozoic sequences west of the Leinster Granite and in the Slievenamon Inlier of southeast Ireland have been palynologically re-investigated. Most of the productive samples yielded sufficient identifiable acritarchs for positive stratigraphical age determinations for several of the formations. The samples also include rare cryptospores, scolecodonts and tubular structures. Previous work in the area west of the Leinster Granite proposed an unbroken succession from Early Ordovician Ribband Group turbidites and volcanics passing up conformably to Early Ordovician to Late Silurian Kilcullen Group. The new palynological data clearly show that the Kilcullen Group in this area is entirely Silurian (Llandovery-early Wenlock) in age, also results obtained from the same group at Slievenamon confirm the previously reported Silurian age. Ordovician acritarchs found in the Kilcullen Group of both study areas are reworked and range in age from late Tremadoc to Llanvirn. The new data reveal a major stratigraphic break between the Ribband Group dated as Early and Middle Ordovician and the Silurian Kilcullen Group. This major break extends some hundreds of kms southwest to the Dingle Peninsula and possibly equates with a similar discontinuity in the Isle of Man to the northeast. This break would thus appear to be a major feature within the northwestern Avalonian margin sequence.     

Early post-mortem calcified Devonian acritarchs as a source of calcispheric structures

Uniquely preserved Late Devonian calcispheres were found in a core of the deep borehole Sosnowiec IG-1 (Upper Silesia, southern Poland). These enigmatic calcareous microfossils are interpreted here as acritarchs that underwent an early post-mortem calcification. Remnants of organic walls preserved in the calcispheres suggest that they represent various acanthomorphic acritarchs, characteristic members of the Palaeozoic marine phytoplankton. Taphonomic analysis combined with the light microscope and scanning electron microscope (SEM) observations of mineral and organic components of the investigated calcispheres suggest that a complex multi-stage process led to calcification of their in vivo non-mineralized acritarch forerunners. The ubiquity of acanthomorphic calcispheres in many Devonian shallow-water limestones is a testimony to little, thus far, documented acritarch crops that must have existed over extensive areas of carbonate-producing epicontinental seas. The scarcity of acritarchs described from Devonian shallow-water limestones may thus represent a taphonomic bias rather than real rarity or absence.     

Palynostratigraphic,palynofacies, organic geochemical and palaeoenvironmental analysis of the Silurian Tanezzuft Formation in the Ghadames Basin of north-west Libya

Two palynofacies associations are documented from the Silurian Tanezzuft Formation in the Ghadames Basin. These are characteristic of the basal ‘Hot Shale’ and the overlying deposits, referred to here as the Cold Shale. The former reflects deposition in distal suboxic anoxic conditions and is dominated by highly oil-prone amorphous organic matter (AOM) typical of deposition in generally anoxic, restricted marine basins. Only a few acritarchs, prasinophyte algae and chitinozoans occur in association: virtually no spores or cryptospores were recorded. Thick-walled prasinophytes are most numerous in this part of the Tanezzuft Formation in both the Ghadames and Murzuq basins, suggesting enhanced surface water productivity. Deposition took place after the melting of the Late Ordovician ice sheets, which led to a major marine transgression. The palynofacies recorded from the overlying ‘Cold Shale’ deposits indicate deposition in distal shelf and basin conditions that were also relatively anoxic. They contain more palynomorphs, especially acritarchs, and generally less AOM. The phytoplankton assemblages are dominated by simple and thin-walled prasinophyte algae (leiospheres), suggesting dysoxic–anoxic conditions. Overall the middle and the upper parts of the Tanezzuft Formation are regarded as being deposited in distal dysoxic–anoxic shelf, distal dysoxic–oxic shelf and distal suboxic–anoxic basin respectively. Because chitinozoans are very rare, age determinations of the samples investigated are based mainly on acritarchs. The Hot Shale is dated as early-mid Rhuddanian (early Llandovery) whereas the rest of the formation is considered to late Rhuddanian–Telychian in age. As documented previously from other samples of the Tanezzuft Formation in both the Ghadames and Murzuq basins, the Hot Shale has a very high TOC content and excellent source potential for liquid hydrocarbons, whereas rest of the formation is less rich in organic matter with larger terrestrial and oxidized components and hence reduced potential for sourcing hydrocarbons (both oil and gas).