Person: Knoll, Andrew
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Publication An Emerging Picture of Neoproterozoic Ocean Chemistry: Insights from the Chuar Group, Grand Canyon, USA
(Elsevier, 2009) Johnston, David; Poulton, Simon W.; Dehler, Carol; Porter, Susannah; Husson, Jon; Canfield, Donald E.; Knoll, AndrewDetailed iron, sulfur and carbon chemistry through the > 742 million year old ChuarGroup reveals a marine basin dominated by anoxic and ferrous iron-rich (ferruginous) bottom waters punctuated, late in the basin's development, by an intrusion of sulfide-rich (euxinic) conditions. The observation that anoxia occurred frequently in even the shallowest of Chuar environments (10s of meters or less) suggests that global atmospheric oxygen levels were significantly lower than today. In contrast, the transition from ferruginous to euxinic subsurface water is interpreted to reflect basinal control—specifically, increased export of organic carbon from surface waters. Low fluxes of organic carbon into subsurface water masses should have been insufficient to deplete oxygen via aerobic respiration, resulting in an oxic oxygen minimum zone (OMZ). Where iron was available, larger organic carbon fluxes should have depleted oxygen and facilitated anaerobic respiration using ferric iron as the oxidant, with iron carbonate as the expected mineralogical signature in basinal shale. Even higher organic fluxes would, in turn, have depleted ferric iron and up-regulated anaerobic respiration by sulfate reduction, reflected in high pyrite abundances. Observations from the ChuarGroup are consistent with these hypotheses, and gain further support from pyrite and sulfate sulfur isotope abundances. In general, Chuar data support the hypothesis that ferruginous subsurface waters returned to the oceans, replacing euxinia, well before the Ediacaran emergence of persistently oxygenated conditions, and even predating the Sturtian glaciation. Moreover, our data suggest that the reprise of ferruginous water masses may relate to widespread rifting during the break-up of Rodinia. This environmental transition, in turn, correlates with both microfossil and biomarker evidence for an expanding eukaryotic presence in the oceans, suggesting a physiologically mediated link among tectonics, environmental chemistry and life in the dynamic Neoproterozoic Earth system.
Publication A Physiologically Explicit Morphospace for Tracheid-based Water Transport in Modern and Extinct Seed Plants
(Paleontological Society, 2010) Wilson, Jonathan P.; Knoll, AndrewWe present a morphometric analysis of water transport cells within a physiologically explicit three-dimensional space. Previous work has shown that cell length, diameter, and pit resistance govern the hydraulic resistance of individual conducting cells; thus, we use these three parameters as axes for our morphospace. We compare living and extinct plants within this space to investigate how patterns of plant conductivity have changed over evolutionary time. Extinct coniferophytes fall within the range of living conifers, despite differences in tracheid-level anatomy. Living cycads, Ginkgo biloba, the Miocene fossil Ginkgo beckii, and extinct cycadeoids overlap with both conifers and vesselless angiosperms. Three Paleozoic seed plants, however, occur in a portion of the morphospace that no living seed plant occupies. Lyginopteris, Callistophyton, and, especially, Medullosa evolved tracheids with high conductivities similar to those of some vessel-bearing angiosperms. Such fossils indicate that extinct seed plants evolved a structural and functional diversity of xylem architectures broader, in some ways, than the range observable in living seed plants.
Publication Phylogenetic, Functional, and Geological Perspectives on Complex Multicellularity
(The MIT Press, 2011) Knoll, Andrew; Hewitt, DavidThis chapter develops a subtle model that integrates environmental and internal factors. It describes the phylogenetic distribution of multicellular organisms in general and complex multicellular life in particular, clarifying the important distinction between the two. This chapter shows that the long apparent lag between the appearance of simple multicellularity in eukaryotes and the radiation of groups with complex multicellular organization has an environmental component that can be associated back to the consequences of life with interior and exterior cells. It suggests that the evolutionary transition from unicells to complex multicellular organisms has several steps.
Publication Lynn Margulis, 1938--2011
(National Academy of Sciences, 2011-12-20) Knoll, AndrewPublication Ediacaran Microfossils from the Ura Formation, Baikal-Patom Uplift, Siberia: Taxonomy and Biostratigraphic Significance
(Paleontological Society, 2011) Sergeev, Vladimir N.; Knoll, Andrew; Vorob, Natalya G.Abundant and diverse microfossils from shales of the uppermost Ura Formation, central Siberia, document early to middle Ediacaran life along the southeastern margin of the Siberian Platform. The Ura Formation is well exposed in a series of sections in the Lena River basin but the best microfossil assemblages come from a locality along the Ura River. Here, the uppermost twenty meters of the formation contain diverse microfossils exceptionally well preserved as organic compressions. Fossils include nearly two dozen morphospecies of large acanthomorphic microfossils attributable to the Ediacaran Complex Acanthomorph Palynoflora (ECAP), a distinctive assemblage known elsewhere only from lower, but not lowermost, to middle Ediacaran rocks. Discovery of ECAP in strata previously considered Mesoproterozoic through Cryogenian confirms inferences from chemostratigraphy, dramatically changing stratigraphic interpretation of sedimentary successions and Proterozoic tectonics on the Siberian Platform. Systematic paleontology is reported for 36 taxa (five described informally) assigned to 23 genera of both eukaryotic and prokaryotic microfossils. One new genus and two new species are proposed: Ancorosphaeridium magnum n. gen. n. sp. and A. minor n. gen. n. sp.
Publication The Deep History of Life
(ASM Press, 2012) Knoll, AndrewThe conventional fossil record is built of hard parts—bones, shells, and decay-resistant organic tissues buried in the sediments that accumulate on floodplains, in lakes, and on the seafloor. In the 1950s, geologists first began the routine application of radioactive decay to problems of geologic age. The geologic record of microbial life is preserved in four distinct ways. First, bacteria and protists leave what we can consider an extension of the conventional fossil record: cell walls and extracellular envelopes preserved directly in sedimentary rocks. Second, microorganisms also leave molecular fossils that complement the record of morphology. Sediments transported across the seafloor interact physically with microbial mat communities, providing a third and distinctly different biological signature in sedimentary rocks. Finally, microbial populations can actually influence the composition of seawater, providing a distinct chemical signature in minerals precipitated from ancient oceans. One can hazard only broad guesses about the biological properties of early microorganisms, but one can make one key statement with confidence: early cells lived without oxygen. The plants and animals so conspicuous in our own world are evolutionary latecomers, intercalated into ecosystems that were already 3 billion years old when sponges first gained a foothold on the seafloor. The author suspects that the correct explanation will not point to physical or biological processes acting alone but, rather, will emphasize the interactions between Earth and life.
Publication Did sulfate availability facilitate the evolutionary expansion of chlorophyll a+c phytoplankton in the oceans?
(Wiley-Blackwell, 2011) Ratti, S.; Knoll, Andrew; Giordano, M.During the Mesozoic Era, dinoflagellates, coccolithophorids and diatoms became prominent primary producers in the oceans, succeeding an earlier biota in which green algae and cyanobacteria had been proportionally more abundant. This transition occurred during an interval marked by increased sulfate concentration in seawater. To test whether increasing sulfate availability facilitated the evolutionary transition in marine phytoplankton, the cyanobacterium Synechococcus sp., the green alga Tetraselmis suecica and three algae containing chlorophyll a+c (the diatom Thalassiosira weissflogii, the dinoflagellate Protoceratium reticulatum and the coccolithophorid Emiliania huxleyi) were grown in media containing 1, 5, 10, 20, or 30 mm SO42−. The cyanobacterium and the green alga showed no growth response to varying [SO42−]. By contrast, the three chlorophyll a+c algae showed improved growth with higher [SO42−], but only up to 10 mm. The chlorophyll a+c algae, but not the green alga or cyanobacterium, also showed lower C:S with higher [SO42−]. When the same experiment was repeated in the presence of a ciliate predator (Euplotes sp.), T. suecica and T. weissflogii increased their specific growth rate in most treatments, whereas the growth rate of Synechococcus sp. was not affected or decreased in the presence of grazers. In a third experiment, T. suecica, T. weissflogii, P. reticulatum and Synechococcus sp. were grown in conditions approximating modern, earlier Paleozoic and Proterozoic seawater. In these treatments, sulfate availability, nitrogen source, metal availability and Pco2 varied. Monospecific cultures exhibited their highest growth rates in the Proterozoic treatment. In mixed culture, T. weissflogii outgrew other species in modern seawater and T. suecica outgrew the others in Paleozoic water. Synechococcus sp. grew best in Proterozoic seawater, but did not outgrow eukaryotic species in any treatment. Collectively, our results suggest that secular increase in seawater [SO42−] may have facilitated the evolutionary expansion of chlorophyll a+c phytoplankton, but probably not to the exclusion of other biological and environmental factors.
Publication Estimating the timing of early eukaryotic diversification with multigene molecular clocks
(Proceedings of the National Academy of Sciences, 2011) Parfrey, L. W.; Lahr, D. J. G.; Knoll, Andrew; Katz, L. A.Although macroscopic plants, animals, and fungi are the most familiar eukaryotes, the bulk of eukaryotic diversity is microbial. Elucidating the timing of diversification among the more than 70 lineages is key to understanding the evolution of eukaryotes. Here, we use taxon-rich multigene data combined with diverse fossils and a relaxed molecular clock framework to estimate the timing of the last common ancestor of extant eukaryotes and the divergence of major clades. Overall, these analyses suggest that the last common ancestor lived between 1866 and 1679 Ma, consistent with the earliest microfossils interpreted with confidence as eukaryotic. During this interval, the Earth's surface differed markedly from today; for example, the oceans were incompletely ventilated, with ferruginous and, after about 1800 Ma, sulfidic water masses commonly lying beneath moderately oxygenated surface waters. Our time estimates also indicate that the major clades of eukaryotes diverged before 1000 Ma, with most or all probably diverging before 1200 Ma. Fossils, however, suggest that diversity within major extant clades expanded later, beginning about 800 Ma, when the oceans began their transition to a more modern chemical state. In combination, paleontological and molecular approaches indicate that long stems preceded diversification in the major eukaryotic lineages.
Publication Preservation of Martian Organic and Environmental Records: Final Report of the Mars Biosignature Working Group
(Mary Ann Liebert Inc, 2011) Summons, Roger E.; Amend, Jan P.; Bish, David; Buick, Roger; Cody, George D.; Des Marais, David; Dromart, Gilles; Eigenbrode, Jennifer L.; Knoll, Andrew; Sumner, Dawn Y.The Mars Science Laboratory (MSL) has an instrument package capable of making measurements of past and present environmental conditions. The data generated may tell us if Mars is, or ever was, able to support life. However, the knowledge of Mars' past history and the geological processes most likely to preserve a record of that history remain sparse and, in some instances, ambiguous. Physical, chemical, and geological processes relevant to biosignature preservation on Earth, especially under conditions early in its history when microbial life predominated, are also imperfectly known. Here, we present the report of a working group chartered by the Co-Chairs of NASA's MSL Project Science Group, John P. Grotzinger and Michael A. Meyer, to review and evaluate potential for biosignature formation and preservation on Mars. Orbital images confirm that layered rocks achieved kilometer-scale thicknesses in some regions of ancient Mars. Clearly, interplays of sedimentation and erosional processes govern present-day exposures, and our understanding of these processes is incomplete. MSL can document and evaluate patterns of stratigraphic development as well as the sources of layered materials and their subsequent diagenesis. It can also document other potential biosignature repositories such as hydrothermal environments. These capabilities offer an unprecedented opportunity to decipher key aspects of the environmental evolution of Mars' early surface and aspects of the diagenetic processes that have operated since that time. Considering the MSL instrument payload package, we identified the following classes of biosignatures as within the MSL detection window: organism morphologies (cells, body fossils, casts), biofabrics (including microbial mats), diagnostic organic molecules, isotopic signatures, evidence of biomineralization and bioalteration, spatial patterns in chemistry, and biogenic gases. Of these, biogenic organic molecules and biogenic atmospheric gases are considered the most definitive and most readily detectable by MSL.
Publication Morphospaces and Databases: Diatom Diversification through Time
(Springer Science + Business Media, 2015) Kotrc, Benjamin; Knoll, AndrewThe diversity of diatom form inspired Art Nouveau designers, an interest renewed by recent advances in biomimetic design. The fossil record provides two windows on the diversification history of diatoms: taxonomic diversity and morphological disparity. Marine planktonic diatom diversity is conventionally interpreted to describe a steep, almost monotonic rise through Cenozoic time. Subsampling methods used to address the associated rise in sampling reveal a more stationary pattern, with peak diversity in the mid-Cenozoic, whether by established methods or a new method (shareholder quorum subsampling, SQS). However, these methods may underestimate diversification if evenness decreases. In order to measure morphological disparity, we constructed an empirical morphospace based on discrete characters. Mean pairwise distance, a disparity metric describing the density of taxa in morphospace, shows little secular change , while convex hull volume, a measure of the extent of occupied morphospace, increases through time. Since we populated the morphospace with occurrence-based data, we can apply subsampling algorithms to these disparity metrics. Mean pairwise distance is largely unaffected, while the increase in occupied volume largely disappears under subsampling. Depending on the metric used, characterizing diatom diversification thus depends upon whether a literal reading of the fossil record or the use of subsampling algorithms is preferred. While this may prompt a reexamination of evolutionary narratives prominently featuring diatom diversification, changes in abundance and silicification may also affect the diatom’s biogeochemical importance. For biologically inspired design, an early exploration of diatom morphospace suggests that fossil forms should be considered alongside extant diatoms.