Ordovician and Silurian sea–water chemistry, sea level, and climate: A synopsis

The Great Ordovician Biodiversification Event (GOBE): The palaeoecological dimension

Rapid and sustained biotic diversification reached its highest levels in the Paleozoic. A prolonged “hot-house” climate through Early Ordovician, cooling through Middle Ordovician and changing to “ice-house” conditions in Late Ordovician, global glaciation, oceanic turnover and mass extinction at end of period, strong fluctuations in eustatic sea level, appearance and diversification of pandemic planktonic graptolites and conodonts important for correlation, moderate to strong benthic faunal provincialism, re-organization and rapid migration of tectonic plates surrounding the Iapetus Ocean and migration of the South Pole from North Africa to central Africa all characterize the Ordovician Period. All seven Ordovician stages have formalized GSSPs.

The Ordovician Period

The early evolutionary history of the chloroplast lineage remains an open question. It is widely accepted that the endosymbiosis that established the chloroplast lineage in eukaryotes can be traced back to a single event, in which a cyanobacterium was incorporated into a protistan host. It is still unclear, however, which Cyanobacteria are most closely related to the chloroplast, when the plastid lineage first evolved, and in what habitats this endosymbiotic event occurred. We present phylogenomic and molecular clock analyses, including data from cyanobacterial and chloroplast genomes using a Bayesian approach, with the aim of estimating the age for the primary endosymbiotic event, the ages of crown groups for photosynthetic eukaryotes, and the independent incorporation of a cyanobacterial endosymbiont by Paulinella. Our analyses include both broad taxon sampling (119 taxa) and 18 fossil calibrations across all Cyanobacteria and photosynthetic eukaryotes. Phylogenomic analyses support the hypothesis that the chloroplast lineage diverged from its closet relative Gloeomargarita, a basal cyanobacterial lineage, ∼2.1 billion y ago (Bya). Our analyses suggest that the Archaeplastida, consisting of glaucophytes, red algae, green algae, and land plants, share a common ancestor that lived ∼1.9 Bya. Whereas crown group Rhodophyta evolved in the Mesoproterozoic Era (1,600–1,000 Mya), crown groups Chlorophyta and Streptophyta began to radiate early in the Neoproterozoic (1,000–542 Mya). Stochastic mapping analyses indicate that the first endosymbiotic event occurred in low-salinity environments. Both red and green algae colonized marine environments early in their histories, with prasinophyte green phytoplankton diversifying 850–650 Mya.

/pdf/early-photosynthetic-eukaryotes-inhabited-low-salinity-3fl45llatv.pdf

Early photosynthetic eukaryotes inhabited low-salinity habitats

The Late Ordovician (Katian-Hirnantian) through earliest Silurian (Rhuddanian) interval was a time of varying climate and sea level, marked by a peak glacial episode in the early-mid-Hirnantian. Synthesis of recently published data permits global correlation of at least two cycles of glacial advance and retreat with a distinct interglacial period that is recognizable in sequence-stratigraphic and chemostratigraphic records in many parts of the world. A period of warming and sea-level rise during the late Katian is marked by the widespread occurrences of oceanic anoxia in paleotropical and subtropical localities, mostly confined to regions of inferred upwelling and semirestricted marine basins. Nitrogen isotope data show that the regions of oceanic anoxia were marked by intense water-column denitrification in which cyanobacteria were the principal source of fixed N. In the overlying peak glacial interval of the Hirnantian, sedimentary successions from localities representing a wide range of water depths and paleolatitudes indicate that anoxia was restricted during the early-mid-Hirnantian. The shift to more positive N isotope values also suggests less intense water-column denitrification. In the overlying late Hirnantian and early Rhuddanian, the distribution of black shales reaches its greatest extent in the studied interval. Localities showing evidence of anoxia are globally spread over all paleolatitudes and water depths for which data are available, indicating a Rhuddanian ocean anoxic event comparable to examples from the Mesozoic. It is accompanied by a return to intensely denitrifying conditions within the water column, as indicated by the shift to negative N isotope values. The two phases of Hirnantian mass extinction coincide with rapid, climate-driven changes in oceanic anoxia. The first extinction occurred at the onset of glaciation and with the loss of anoxic conditions at the end of the Katian. The second extinction occurred at the demise of glaciation and coincided with the return of anoxic conditions during the late Hirnantian–early Rhuddanian. Integration of our N isotope data with graptolite biodiversity records suggests that the extinctions were profoundly influenced by changes occurring at the base of the marine food web, i.e., redox-driven changes in nutrient cycling and primary producer communities.

Environmental changes in the Late Ordovician–early Silurian: Review and new insights from black shales and nitrogen isotopes

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 delta C-13(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.

The Ordovician Biodiversification: revolution in the oceanic trophic chain

Klug, C., Kroger, B., Kiessling, W., Mullins, G.L., Servais, T., Frýda, J., Korn, D. & Turner, S. 2009: The Devonian nekton revolution. Lethaia, 10.1111/j.1502-3931.2009.00206.x

Traditional analyses of Early Phanerozoic marine diversity at the genus level show an explosive radiation of marine life until the Late Ordovician, followed by a phase of erratic decline continuing until the end of the Palaeozoic, whereas a more recent analysis extends the duration of this early radiation into the Devonian. This catch-all approach hides an evolutionary and ecological key event long after the Ordovician radiation: the rapid occupation of the free water column by animals during the Devonian. Here, we explore the timing of the occupation of the water column in the Palaeozoic and test the hypothesis that ecological escalation led to fundamental evolutionary changes in the mid-Palaeozoic marine water column. According to our analyses, demersal and nektonic modes of life were probably initially driven by competition in the diversity-saturated benthic habitats together with the availability of abundant planktonic food. Escalatory feedback then promoted the rapid rise of nekton in the Devonian as suggested by the sequence and tempo of water-column occupation. □Devonian, diversity, ecology, food webs, nekton, plankton, radiation.

/pdf/the-devonian-nekton-revolution-2m1qdxk3i9.pdf

The Devonian nekton revolution

Numerous environmental factors as well as oceanic circulation patterns and geographic constraints all contribute to the abundance, distribution, and diversity of present-day marine phytoplankton assemblages. These same factors presumably affected the Paleozoic marine phytoplankton, which was dominated by organic-walled acritarchs and prasinophytes. During the Late Silurian (Gorstian, Ludfordian, and Přidoli) and earliest Devonian (Lochkovian), important paleogeographic, paleooceanographic, and geochemical changes were occurring as well as major compositional changes and diversity fluctuations in the marine organic-walled phytoplankton. Innovative morphologies appeared during the Late Silurian, in both low and high latitude assemblages, but with significant quantitative differences. This was followed by a turnover in assemblage composition during the Silurian/Devonian transition, and an initial radiation of new acritarch and prasinophyte taxa in the Early Devonian. Observed changes in total phytop...

Global patterns of organic-walled phytoplankton biodiversity during the late silurian to earliest devonian

The diversity of the Carboniferous phytoplankton

Diverse and abundant assemblages of chitinozoans allow the recognition of the upper part of the Margachitina margaritana Biozone in the uppermost Llandovery and lowermost Wenlock series of the Hughley Brook section. The Cingulochitina bouniensis and Salopochitina bella biozones are identified in the lower part of the Buildwas Formation, Wenlock Series. The chitinozoan data indicate that the base of the Wenlock Series most probably correlates with a level in the upper centrifugus or lower murchisoni graptolite biozones. Chitinozoans also indicate that the base of the riccartonensis graptolite Biozone may occur within the Buildwas Formation and not the overlying Coalbrookdale Formation.

Gary L. Mullins

Papers

The Ordovician Biodiversification: revolution in the oceanic trophic chain

The Devonian nekton revolution

Global patterns of organic-walled phytoplankton biodiversity during the late silurian to earliest devonian

The diversity of the Carboniferous phytoplankton

Chitinozoan biostratigraphy of the basal Wenlock Series (Silurian) Global Stratotype Section and Point