Academic Appointments

Honors & Awards

  • Explorer, National Geographic (2017)
  • Ocean Sciences Fellowship, Alfred P. Sloan Foundation (2017)
  • Pre-tenure award, GSA Geobiology and Geomicrobiology Division (2016)
  • NAI Postdoctoral Fellowship, NASA Astrobiology Institute (2012-2014)
  • Geobiology Fellowship, Agouron Institute (2010-2012)

Boards, Advisory Committees, Professional Organizations

  • Editorial Board, Palaios (2017 - Present)
  • Editorial Advisory Board, Geobiology (2016 - Present)

Current Research and Scholarly Interests

The research interests in the Sperling Lab are Earth history and the evolution of life, and the interactions between the biosphere and the geosphere. As such this research can generally be considered paleontology, insofar as paleontology encompasses all aspects of the history of life.

Consequently, we define our research agenda by the questions we are interested in, rather than the tools used. This research incorporates multiple lines of evidence, and multiple tools, to investigate questions in the history of life. These lines of evidence include fossil data, molecular phylogenetics, sedimentary geochemistry, and developmental and ecological data from modern organisms. Ultimately, the goal is to link environmental change with organismal and ecological response through the lens of physiology.

Our field research takes place all over the world--current areas include:

-NW Canada (Yukon and Northwest Territories): Research has been conducted on the early Neoproterozoic Fifteenmile Group, Cryogenian and Ediacaran Windermere Supergroup, and on the Ordovician-Devonian Road River Group in the southern Richardson Mountains
-Southern Canadian Cordillera: Work here has focused on the early Cambrian Mural Formation and its soft-bodied fauna.
-England and Wales: Cambrian-Silurian successions in the Welsh Basin
-Namibia: Ediacaran Nama Group
-Upwelling zones: We study the oxygen minimum zone offshore California as an analogue for ancient low-oxygen oceans.

2023-24 Courses

Stanford Advisees

All Publications

  • The relationship between total organic carbon and bottom water redox state in North American black shales PALAEOGEOGRAPHY PALAEOCLIMATOLOGY PALAEOECOLOGY Ritzer, S. R., Schoepfer, S., Bussian, B., Farrell, U. C., Fraser, T., Henderson, C. M., Kang, J., Mwinde, C. N., Patch, A., Sperling, E. A. 2024; 649
  • A shift in redox conditions near the Ediacaran/Cambrian transition and its possible influence on early animal evolution, Corumb4Group, Brazil GEOSCIENCE FRONTIERS Caxito, F. A., Sperling, E., Fazio, G., Adorno, R., Denezine, M., Do Carmo, D., Giorgioni, M., Uhlein, G. J., Sial, A. N. 2024; 15 (4)
  • Lithium isotopic constraints on the evolution of continental clay mineral factory and marine oxygenation in the earliest Paleozoic Era. Science advances Wei, G. Y., Zhao, M., Sperling, E. A., Gaines, R. R., Kalderon-Asael, B., Shen, J., Li, C., Zhang, F., Li, G., Zhou, C., Cai, C., Chen, D., Xiao, K. Q., Jiang, L., Ling, H. F., Planavsky, N. J., Tarhan, L. G. 2024; 10 (13): eadk2152


    The evolution of oxygen cycles on Earth's surface has been regulated by the balance between molecular oxygen production and consumption. The Neoproterozoic-Paleozoic transition likely marks the second rise in atmospheric and oceanic oxygen levels, widely attributed to enhanced burial of organic carbon. However, it remains disputed how marine organic carbon production and burial respond to global environmental changes and whether these feedbacks trigger global oxygenation during this interval. Here, we report a large lithium isotopic and elemental dataset from marine mudstones spanning the upper Neoproterozoic to middle Cambrian [~660 million years ago (Ma) to 500 Ma]. These data indicate a dramatic increase in continental clay formation after ~525 Ma, likely linked to secular changes in global climate and compositions of the continental crust. Using a global biogeochemical model, we suggest that intensified continental weathering and clay delivery to the oceans could have notably increased the burial efficiency of organic carbon and facilitated greater oxygen accumulation in the earliest Paleozoic oceans.

    View details for DOI 10.1126/sciadv.adk2152

    View details for PubMedID 38552018

  • Deep-water first occurrences of Ediacara biota prior to the Shuram carbon isotope excursion in the Wernecke Mountains, Yukon, Canada. Geobiology Boag, T. H., Busch, J. F., Gooley, J. T., Strauss, J. V., Sperling, E. A. 2024; 22 (3): e12597


    Ediacara-type macrofossils appear as early as ~575 Ma in deep-water facies of the Drook Formation of the Avalon Peninsula, Newfoundland, and the Nadaleen Formation of Yukon and Northwest Territories, Canada. Our ability to assess whether a deep-water origination of the Ediacara biota is a genuine reflection of evolutionary succession, an artifact of an incomplete stratigraphic record, or a bathymetrically controlled biotope is limited by a lack of geochronological constraints and detailed shelf-to-slope transects of Ediacaran continental margins. The Ediacaran Rackla Group of the Wernecke Mountains, NW Canada, represents an ideal shelf-to-slope depositional system to understand the spatiotemporal and environmental context of Ediacara-type organisms' stratigraphic occurrence. New sedimentological and paleontological data presented herein from the Wernecke Mountains establish a stratigraphic framework relating shelfal strata in the Goz/Corn Creek area to lower slope deposits in the Nadaleen River area. We report new discoveries of numerous Aspidella hold-fast discs, indicative of frondose Ediacara organisms, from deep-water slope deposits of the Nadaleen Formation stratigraphically below the Shuram carbon isotope excursion (CIE) in the Nadaleen River area. Such fossils are notably absent in coeval shallow-water strata in the Goz/Corn Creek region despite appropriate facies for potential preservation. The presence of pre-Shuram CIE Ediacara-type fossils occurring only in deep-water facies within a basin that has equivalent well-preserved shallow-water facies provides the first stratigraphic paleobiological support for a deep-water origination of the Ediacara biota. In contrast, new occurrences of Ediacara-type fossils (including juvenile fronds, Beltanelliformis, Aspidella, annulated tubes, and multiple ichnotaxa) are found above the Shuram CIE in both deep- and shallow-water deposits of the Blueflower Formation. Given existing age constraints on the Shuram CIE, it appears that Ediacaran organisms may have originated in the deeper ocean and lived there for up to ~15 million years before migrating into shelfal environments in the terminal Ediacaran. This indicates unique ecophysiological constraints likely shaped the initial habitat preference and later environmental expansion of the Ediacara biota.

    View details for DOI 10.1111/gbi.12597

    View details for PubMedID 38700422

  • Constraining the oxygen requirements for modern microbial eukaryote diversity. Proceedings of the National Academy of Sciences of the United States of America Mills, D. B., Simister, R. L., Sehein, T. R., Hallam, S. J., Sperling, E. A., Crowe, S. A. 2024; 121 (2): e2303754120


    Eukaryotes originated prior to the establishment of modern marine oxygen (O2) levels. According to the body fossil and lipid biomarker records, modern (crown) microbial eukaryote lineages began diversifying in the ocean no later than ~800 Ma. While it has long been predicted that increasing atmospheric O2 levels facilitated the early diversification of microbial eukaryotes, the O2 levels needed to permit this diversification remain unconstrained. Using time-resolved geochemical parameter and gene sequence information from a model marine oxygen minimum zone spanning a range of dissolved O2 levels and redox states, we show that microbial eukaryote taxonomic richness and phylogenetic diversity remain the same until O2 declines to around 2 to 3% of present atmospheric levels, below which these diversity metrics become significantly reduced. Our observations suggest that increasing O2 would have only directly promoted early crown-eukaryote diversity if atmospheric O2 was below 2 to 3% of modern levels when crown-eukaryotes originated and then later met or surpassed this range as crown-eukaryotes diversified. If atmospheric O2 was already consistently at or above 2 to 3% of modern levels by the time that crown-eukaryotes originated, then the subsequent diversification of modern microbial eukaryotes was not directly driven by atmospheric oxygenation.

    View details for DOI 10.1073/pnas.2303754120

    View details for PubMedID 38165897

  • Thermal optima in the hypoxia tolerance of marine ectotherms: Physiological causes and biogeographic consequences. PLoS biology Endress, M. A., Penn, J. L., Boag, T. H., Burford, B. P., Sperling, E. A., Deutsch, C. A. 2024; 22 (1): e3002443


    The minimum O2 needed to fuel the demand of aquatic animals is commonly observed to increase with temperature, driven by accelerating metabolism. However, recent measurements of critical O2 thresholds ("Pcrit") reveal more complex patterns, including those with a minimum at an intermediate thermal "optimum". To discern the prevalence, physiological drivers, and biogeographic manifestations of such curves, we analyze new experimental and biogeographic data using a general dynamic model of aquatic water breathers. The model simulates the transfer of oxygen from ambient water through a boundary layer and into animal tissues driven by temperature-dependent rates of metabolism, diffusive gas exchange, and ventilatory and circulatory systems with O2-protein binding. We find that a thermal optimum in Pcrit can arise even when all physiological rates increase steadily with temperature. This occurs when O2 supply at low temperatures is limited by a process that is more temperature sensitive than metabolism, but becomes limited by a less sensitive process at warmer temperatures. Analysis of published species respiratory traits suggests that this scenario is not uncommon in marine biota, with ventilation and circulation limiting supply under cold conditions and diffusion limiting supply at high temperatures. Using occurrence data, we show that species with these physiological traits inhabit lowest O2 waters near the optimal temperature for hypoxia tolerance and are restricted to higher O2 at temperatures above and below this optimum. Our results imply that hypoxia tolerance can decline under both cold and warm conditions and thus may influence both poleward and equatorward species range limits.

    View details for DOI 10.1371/journal.pbio.3002443

    View details for PubMedID 38227580

    View details for PubMedCentralID PMC10790991

  • Oxygen availability and body mass modulate ectotherm responses to ocean warming. Nature communications Duncan, M. I., Micheli, F., Boag, T. H., Marquez, J. A., Deres, H., Deutsch, C. A., Sperling, E. A. 2023; 14 (1): 3811


    In an ocean that is rapidly warming and losing oxygen, accurate forecasting of species' responses must consider how this environmental change affects fundamental aspects of their physiology. Here, we develop an absolute metabolic index (ΦA) that quantifies how ocean temperature, dissolved oxygen and organismal mass interact to constrain the total oxygen budget an organism can use to fuel sustainable levels of aerobic metabolism. We calibrate species-specific parameters of ΦA with physiological measurements for red abalone (Haliotis rufescens) and purple urchin (Strongylocentrotus purpuratus). ΦA models highlight that the temperature where oxygen supply is greatest shifts cooler when water loses oxygen or organisms grow larger, providing a mechanistic explanation for observed thermal preference patterns. Viable habitat forecasts are disproportionally deleterious for red abalone, revealing how species-specific physiologies modulate the intensity of a common climate signal, captured in the newly developed ΦA framework.

    View details for DOI 10.1038/s41467-023-39438-w

    View details for PubMedID 37369654

    View details for PubMedCentralID PMC10300008

  • Species of Dickinsonia Sprigg from the Ediacaran of South Australia PALAEONTOLOGY Evans, S. D., Hunt, G., Gehling, J. G., Sperling, E. A., Droser, M. L. 2023; 66 (1)

    View details for DOI 10.1111/pala.12635

    View details for Web of Science ID 000928749900001

  • Integrated Litho-, Chemo- and Sequence Stratigraphy of the Ediacaran Gametrail Formation Across a Shelf-Slope Transect in the Wernecke Mountains, Yukon, Canada AMERICAN JOURNAL OF SCIENCE Busch, J. F., Boag, T. H., Sperling, E. A., Rooney, A. D., Feng, X., Moynihan, D. P., Strauss, J. V. 2023; 323

    View details for DOI 10.2475/001c.74874

    View details for Web of Science ID 001079515700001

  • Breathless through Time: Oxygen and Animals across Earth's History BIOLOGICAL BULLETIN Sperling, E. A., Boag, T. H., Duncan, M. I., Endriga, C. R., Marquez, J., Mills, D. B., Monarrez, P. M., Sclafani, J. A., Stockey, R. G., Payne, J. L. 2022

    View details for DOI 10.1086/721754

    View details for Web of Science ID 000864473500001

  • Integrative Approaches to Understanding Organismal Responses to Aquatic Deoxygenation BIOLOGICAL BULLETIN Woods, H., Moran, A. L., Atkinson, D., Audzijonyte, A., Berenbrink, M., Borges, F. O., Burnett, K. G., Burnett, L. E., Coates, C. J., Collin, R., Costa-Paiva, E. M., Duncan, M. I., Ern, R., Laetz, E. J., Levin, L. A., Lindmark, M., Lucey, N. M., McCormick, L. R., Pierson, J. J., Rosa, R., Roman, M. R., Sampaio, E., Schulte, P. M., Sperling, E. A., Walczynska, A., Verberk, W. P. 2022: 85-103


    AbstractOxygen bioavailability is declining in aquatic systems worldwide as a result of climate change and other anthropogenic stressors. For aquatic organisms, the consequences are poorly known but are likely to reflect both direct effects of declining oxygen bioavailability and interactions between oxygen and other stressors, including two-warming and acidification-that have received substantial attention in recent decades and that typically accompany oxygen changes. Drawing on the collected papers in this symposium volume ("An Oxygen Perspective on Climate Change"), we outline the causes and consequences of declining oxygen bioavailability. First, we discuss the scope of natural and predicted anthropogenic changes in aquatic oxygen levels. Although modern organisms are the result of long evolutionary histories during which they were exposed to natural oxygen regimes, anthropogenic change is now exposing them to more extreme conditions and novel combinations of low oxygen with other stressors. Second, we identify behavioral and physiological mechanisms that underlie the interactive effects of oxygen with other stressors, and we assess the range of potential organismal responses to oxygen limitation that occur across levels of biological organization and over multiple timescales. We argue that metabolism and energetics provide a powerful and unifying framework for understanding organism-oxygen interactions. Third, we conclude by outlining a set of approaches for maximizing the effectiveness of future work, including focusing on long-term experiments using biologically realistic variation in experimental factors and taking truly cross-disciplinary and integrative approaches to understanding and predicting future effects.

    View details for DOI 10.1086/722899

    View details for Web of Science ID 000898698100001

    View details for PubMedID 36548975

  • Mesoproterozoic surface oxygenation accompanied major sedimentary manganese deposition at 1.4 and 1.1 Ga. Geobiology Spinks, S. C., Sperling, E. A., Thorne, R. L., LaFountain, F., White, A. J., Armstrong, J., Woltering, M., Tyler, I. M. 2022


    Manganese (Mn) oxidation in marine environments requires oxygen (O2 ) or other reactive oxygen species in the water column, and widespread Mn oxide deposition in ancient sedimentary rocks has long been used as a proxy for oxidation. The oxygenation of Earth's atmosphere and oceans across the Archean-Proterozoic boundary are associated with massive Mn deposits, whereas the interval from 1.8-1.0 Ga is generally believed to be a time of low atmospheric oxygen with an apparent hiatus in sedimentary Mn deposition. Here, we report geochemical and mineralogical analyses from 1.1 Ga manganiferous marine-shelf siltstones from the Bangemall Supergroup, Western Australia, which underlie recently discovered economically significant manganese deposits. Layers bearing Mn carbonate microspheres, comparable with major global Mn deposits, reveal that intense periods of sedimentary Mn deposition occurred in the late Mesoproterozoic. Iron geochemical data suggest anoxic-ferruginous seafloor conditions at the onset of Mn deposition, followed by oxic conditions in the water column as Mn deposition persisted and eventually ceased. These data imply there was spatially widespread surface oxygenation ~1.1 Ga with sufficiently oxic conditions in shelf environments to oxidize marine Mn(II). Comparable large stratiform Mn carbonate deposits also occur in ~1.4 Ga marine siltstones hosted in underlying sedimentary units. These deposits are greater or at least commensurate in scale (tonnage) to those that followed the major oxygenation transitions from the Neoproterozoic. Such a period of sedimentary manganogenesis is inconsistent with a model of persistently low O2 throughout the entirety of the Mesoproterozoic and provides robust evidence for dynamic redox changes in the mid to late Mesoproterozoic.

    View details for DOI 10.1111/gbi.12524

    View details for PubMedID 36168296

  • Eukaryogenesis and oxygen in Earth history. Nature ecology & evolution Mills, D. B., Boyle, R. A., Daines, S. J., Sperling, E. A., Pisani, D., Donoghue, P. C., Lenton, T. M. 2022


    The endosymbiotic origin of mitochondria during eukaryogenesis has long been viewed as an adaptive response to the oxygenation of Earth's surface environment, presuming a fundamentally aerobic lifestyle for the free-living bacterial ancestors of mitochondria. This oxygen-centric view has been robustly challenged by recent advances in the Earth and life sciences. While the permanent oxygenation of the atmosphere above trace concentrations is now thought to have occurred 2.2 billion years ago, large parts of the deep ocean remained anoxic until less than 0.5 billion years ago. Neither fossils nor molecular clocks correlate the origin of mitochondria, or eukaryogenesis more broadly, to either of these planetary redox transitions. Instead, mitochondria-bearing eukaryotes are consistently dated to between these two oxygenation events, during an interval of pervasive deep-sea anoxia and variable surface-water oxygenation. The discovery and cultivation of the Asgard archaea has reinforced metabolic evidence that eukaryogenesis was initially mediated by syntrophic H2 exchange between an archaeal host and an alpha-proteobacterial symbiont living under anoxia. Together, these results temporally, spatially and metabolically decouple the earliest stages of eukaryogenesis from the oxygen content of the surface ocean and atmosphere. Rather than reflecting the ancestral metabolic state, obligate aerobiosis in eukaryotes is most probably derived, having only become globally widespread over the past 1 billion years as atmospheric oxygen approached modern levels.

    View details for DOI 10.1038/s41559-022-01733-y

    View details for PubMedID 35449457

  • A prolonged, two-step oxygenation of Earth's early atmosphere: Support from confidence intervals GEOLOGY Hodgskiss, M. W., Sperling, E. A. 2022; 50 (2): 158-162

    View details for DOI 10.1130/G49385.1

    View details for Web of Science ID 000749789600006

  • Marine sponges in the once and future ocean. Global change biology Mills, D. B., Sperling, E. A. 1800

    View details for DOI 10.1111/gcb.16053

    View details for PubMedID 34932850

  • Decreasing Phanerozoic extinction intensity as a consequence of Earth surface oxygenation and metazoan ecophysiology PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA Stockey, R. G., Pohl, A., Ridgwell, A., Finnegan, S., Sperling, E. A. 2021; 118 (41)
  • Decreasing Phanerozoic extinction intensity as a consequence of Earth surface oxygenation and metazoan ecophysiology. Proceedings of the National Academy of Sciences of the United States of America Stockey, R. G., Pohl, A., Ridgwell, A., Finnegan, S., Sperling, E. A. 2021; 118 (41)


    The decline in background extinction rates of marine animals through geologic time is an established but unexplained feature of the Phanerozoic fossil record. There is also growing consensus that the ocean and atmosphere did not become oxygenated to near-modern levels until the mid-Paleozoic, coinciding with the onset of generally lower extinction rates. Physiological theory provides us with a possible causal link between these two observations-predicting that the synergistic impacts of oxygen and temperature on aerobic respiration would have made marine animals more vulnerable to ocean warming events during periods of limited surface oxygenation. Here, we evaluate the hypothesis that changes in surface oxygenation exerted a first-order control on extinction rates through the Phanerozoic using a combined Earth system and ecophysiological modeling approach. We find that although continental configuration, the efficiency of the biological carbon pump in the ocean, and initial climate state all impact the magnitude of modeled biodiversity loss across simulated warming events, atmospheric oxygen is the dominant predictor of extinction vulnerability, with metabolic habitat viability and global ecophysiotype extinction exhibiting inflection points around 40% of present atmospheric oxygen. Given this is the broad upper limit for estimates of early Paleozoic oxygen levels, our results are consistent with the relative frequency of high-magnitude extinction events (particularly those not included in the canonical big five mass extinctions) early in the Phanerozoic being a direct consequence of limited early Paleozoic oxygenation and temperature-dependent hypoxia responses.

    View details for DOI 10.1073/pnas.2101900118

    View details for PubMedID 34607946

  • The Sedimentary Geochemistry and Paleoenvironments Project. Geobiology Farrell, U. C., Samawi, R., Anjanappa, S., Klykov, R., Adeboye, O. O., Agic, H., Ahm, A. C., Boag, T. H., Bowyer, F., Brocks, J. J., Brunoir, T. N., Canfield, D. E., Chen, X., Cheng, M., Clarkson, M. O., Cole, D. B., Cordie, D. R., Crockford, P. W., Cui, H., Dahl, T. W., Mouro, L. D., Dewing, K., Dornbos, S. Q., Drabon, N., Dumoulin, J. A., Emmings, J. F., Endriga, C. R., Fraser, T. A., Gaines, R. R., Gaschnig, R. M., Gibson, T. M., Gilleaudeau, G. J., Gill, B. C., Goldberg, K., Guilbaud, R., Halverson, G. P., Hammarlund, E. U., Hantsoo, K. G., Henderson, M. A., Hodgskiss, M. S., Horner, T. J., Husson, J. M., Johnson, B., Kabanov, P., Brenhin Keller, C., Kimmig, J., Kipp, M. A., Knoll, A. H., Kreitsmann, T., Kunzmann, M., Kurzweil, F., LeRoy, M. A., Li, C., Lipp, A. G., Loydell, D. K., Lu, X., Macdonald, F. A., Magnall, J. M., Mand, K., Mehra, A., Melchin, M. J., Miller, A. J., Mills, N. T., Mwinde, C. N., O'Connell, B., Och, L. M., Ossa Ossa, F., Pages, A., Paiste, K., Partin, C. A., Peters, S. E., Petrov, P., Playter, T. L., Plaza-Torres, S., Porter, S. M., Poulton, S. W., Pruss, S. B., Richoz, S., Ritzer, S. R., Rooney, A. D., Sahoo, S. K., Schoepfer, S. D., Sclafani, J. A., Shen, Y., Shorttle, O., Slotznick, S. P., Smith, E. F., Spinks, S., Stockey, R. G., Strauss, J. V., Stueken, E. E., Tecklenburg, S., Thomson, D., Tosca, N. J., Uhlein, G. J., Vizcaino, M. N., Wang, H., White, T., Wilby, P. R., Woltz, C. R., Wood, R. A., Xiang, L., Yurchenko, I. A., Zhang, T., Planavsky, N. J., Lau, K. V., Johnston, D. T., Sperling, E. A. 2021

    View details for DOI 10.1111/gbi.12462

    View details for PubMedID 34219351

  • A long-term record of early to mid-Paleozoic marine redox change. Science advances Sperling, E. A., Melchin, M. J., Fraser, T., Stockey, R. G., Farrell, U. C., Bhajan, L., Brunoir, T. N., Cole, D. B., Gill, B. C., Lenz, A., Loydell, D. K., Malinowski, J., Miller, A. J., Plaza-Torres, S., Bock, B., Rooney, A. D., Tecklenburg, S. A., Vogel, J. M., Planavsky, N. J., Strauss, J. V. 2021; 7 (28)


    The extent to which Paleozoic oceans differed from Neoproterozoic oceans and the causal relationship between biological evolution and changing environmental conditions are heavily debated. Here, we report a nearly continuous record of seafloor redox change from the deep-water upper Cambrian to Middle Devonian Road River Group of Yukon, Canada. Bottom waters were largely anoxic in the Richardson trough during the entirety of Road River Group deposition, while independent evidence from iron speciation and Mo/U ratios show that the biogeochemical nature of anoxia changed through time. Both in Yukon and globally, Ordovician through Early Devonian anoxic waters were broadly ferruginous (nonsulfidic), with a transition toward more euxinic (sulfidic) conditions in the mid-Early Devonian (Pragian), coincident with the early diversification of vascular plants and disappearance of graptolites. This ~80-million-year interval of the Paleozoic characterized by widespread ferruginous bottom waters represents a persistence of Neoproterozoic-like marine redox conditions well into the Phanerozoic.

    View details for DOI 10.1126/sciadv.abf4382

    View details for PubMedID 34233874

  • Isotopic analyses of Ordovician-Silurian siliceous skeletons indicate silica-depleted Paleozoic oceans. Geobiology Trower, E. J., Strauss, J. V., Sperling, E. A., Fischer, W. W. 2021


    The Phanerozoic Eon marked a major transition from marine silica deposition exclusively via abiotic pathways to a system dominated by biogenic silica sedimentation. For decades, prevailing ideas predicted this abiotic-to-biogenic transition were marked by a significant decrease in the concentration of dissolved silica in seawater; however, due to the lower perceived abundance and uptake affinity of sponges and radiolarians relative to diatoms, marine dissolved silica is thought to have remained elevated above modern values until the Cenozoic radiation of diatoms. Studies of modern marine silica biomineralizers demonstrated that the Si isotope ratios (delta30 Si) of sponge spicules and planktonic silica biominerals produced by diatoms or radiolarians can be applied as quantitative proxies for past seawater dissolved silica concentrations due to differences in Si isotope fractionations among these organisms. We undertook 446 ion microprobe analyses of delta30 Si and delta18 O of sponge spicules and radiolarians from Ordovician-Silurian chert deposits of the Mount Hare Formation in Yukon, Canada. These isotopic data showed that sponges living in marine slope and basinal environments displayed small Si isotope fractionations relative to coeval radiolarians. By constructing a mathematical model of the major fluxes and reservoirs in the marine silica cycle and the physiology of silica biomineralization, we found that the concentration of dissolved silica in seawater was less than ~150muM during early Paleozoic time-a value that is significantly lower than previous estimates. We posit that the topology of the early Paleozoic marine silica cycle resembled that of modern oceans much more closely than previously assumed.

    View details for DOI 10.1111/gbi.12449

    View details for PubMedID 34002455

  • Variable redox conditions as an evolutionary driver? A multi-basin comparison of redox in the middle and later Cambrian oceans (Drumian-Paibian) PALAEOGEOGRAPHY PALAEOCLIMATOLOGY PALAEOECOLOGY LeRoy, M. A., Gill, B. C., Sperling, E. A., McKenzie, N., Park, T. S. 2021; 566
  • Thallium isotope ratios in shales from South China and northwestern Canada suggest widespread O-2 accumulation in marine bottom waters was an uncommon occurrence during the Ediacaran Period CHEMICAL GEOLOGY Ostrander, C. M., Owens, J. D., Nielsen, S. G., Lyons, T. W., Shu, Y., Chen, X., Sperling, E. A., Jiang, G., Johnston, D. T., Sahoo, S. K., Anbar, A. D. 2020; 557
  • Redox and paleoenvironmental conditions of the Devonian-Carboniferous Sappington Formation, southwestern Montana, and comparison to the Bakken Formation, Williston Basin PALAEOGEOGRAPHY PALAEOCLIMATOLOGY PALAEOECOLOGY Browne, T. N., Hofmann, M. H., Malkowski, M. A., Wei, J., Sperling, E. A. 2020; 560
  • The Road River Group of northern Yukon, Canada: early Paleozoic deep-water sedimentation within the Great American Carbonate Bank CANADIAN JOURNAL OF EARTH SCIENCES Strauss, J., Fraser, T., Melchin, M. J., Allen, T. J., Malinowski, J., Feng, X., Taylor, J. F., Day, J., Gill, B. C., Sperling, E. A. 2020; 57 (10): 1193–1219
  • Mesoproterozoic paleo-redox changes during 1500-1400 Ma in the Yanshan Basin, North China PRECAMBRIAN RESEARCH Chen, X., Li, M., Sperling, E. A., Zhang, T., Zong, K., Liu, Y., Shen, Y. 2020; 347
  • Extending the record of the Lomagundi-Jatuli carbon isotope excursion in the Labrador Trough, Canada CANADIAN JOURNAL OF EARTH SCIENCES Hodgskiss, M. W., Lamothe, K. G., Halverson, G. P., Sperling, E. A. 2020; 57 (9): 1089–1102
  • A high-TOC shale in a low productivity world: The late Mesoproterozoic Arctic Bay Formation, Nunavut EARTH AND PLANETARY SCIENCE LETTERS Hodgskiss, M. W., Sansjofre, P., Kunzmann, M., Sperling, E. A., Cole, D. B., Crockford, P. W., Gibson, T. M., Halverson, G. R. 2020; 544
  • Uranium Isotope Fractionation in Non-sulfidic Anoxic Settings and the Global Uranium Isotope Mass Balance GLOBAL BIOGEOCHEMICAL CYCLES Cole, D. B., Planavsky, N. J., Longley, M., Boening, P., Wilkes, D., Wang, X., Swanner, E. D., Wittkop, C., Loydell, D. K., Busigny, V., Knudsen, A. C., Sperling, E. A. 2020; 34 (8)
  • Calibrating the coevolution of Ediacaran life and environment. Proceedings of the National Academy of Sciences of the United States of America Rooney, A. D., Cantine, M. D., Bergmann, K. D., Gomez-Perez, I., Al Baloushi, B., Boag, T. H., Busch, J. F., Sperling, E. A., Strauss, J. V. 2020


    The rise of animals occurred during an interval of Earth history that witnessed dynamic marine redox conditions, potentially rapid plate motions, and uniquely large perturbations to global biogeochemical cycles. The largest of these perturbations, the Shuram carbon isotope excursion, has been invoked as a driving mechanism for Ediacaran environmental change, possibly linked with evolutionary innovation or extinction. However, there are a number of controversies surrounding the Shuram, including its timing, duration, and role in the concomitant biological and biogeochemical upheavals. Here we present radioisotopic dates bracketing the Shuram on two separate paleocontinents; our results are consistent with a global and synchronous event between 574.0 ± 4.7 and 567.3 ± 3.0 Ma. These dates support the interpretation that the Shuram is a primary and synchronous event postdating the Gaskiers glaciation. In addition, our Re-Os ages suggest that the appearance of Ediacaran macrofossils in northwestern Canada is identical, within uncertainty, to similar macrofossils from the Conception Group of Newfoundland, highlighting the coeval appearance of macroscopic metazoans across two paleocontinents. Our temporal framework for the terminal Proterozoic is a critical step for testing hypotheses related to extreme carbon isotope excursions and their role in the evolution of complex life.

    View details for DOI 10.1073/pnas.2002918117

    View details for PubMedID 32632000


    View details for DOI 10.1002/spp2.1313

    View details for Web of Science ID 000542124000001

  • Ediacaran reorganization of the marine phosphorus cycle. Proceedings of the National Academy of Sciences of the United States of America Laakso, T. A., Sperling, E. A., Johnston, D. T., Knoll, A. H. 2020


    The Ediacaran Period (635 to 541 Ma) marks the global transition to a more productive biosphere, evidenced by increased availability of food and oxidants, the appearance of macroscopic animals, significant populations of eukaryotic phytoplankton, and the onset of massive phosphorite deposition. We propose this entire suite of changes results from an increase in the size of the deep-water marine phosphorus reservoir, associated with rising sulfate concentrations and increased remineralization of organic P by sulfate-reducing bacteria. Simple mass balance calculations, constrained by modern anoxic basins, suggest that deep-water phosphate concentrations may have increased by an order of magnitude without any increase in the rate of P input from the continents. Strikingly, despite a major shift in phosphorite deposition, a new compilation of the phosphorus content of Neoproterozoic and early Paleozoic shows little secular change in median values, supporting the view that changes in remineralization and not erosional P fluxes were the principal drivers of observed shifts in phosphorite accumulation. The trigger for these changes may have been transient Neoproterozoic weathering events whose biogeochemical consequences were sustained by a set of positive feedbacks, mediated by the oxygen and sulfur cycles, that led to permanent state change in biogeochemical cycling, primary production, and biological diversity by the end of the Ediacaran Period.

    View details for DOI 10.1073/pnas.1916738117

    View details for PubMedID 32424088

  • Persistent global marine euxinia in the early Silurian. Nature communications Stockey, R. G., Cole, D. B., Planavsky, N. J., Loydell, D. K., Fryda, J., Sperling, E. A. 2020; 11 (1): 1804


    The second pulse of the Late Ordovician mass extinction occurred around the Hirnantian-Rhuddanian boundary (~444Ma) and has been correlated with expanded marine anoxia lasting into the earliest Silurian. Characterization of the Hirnantian ocean anoxic event has focused on the onset of anoxia, with global reconstructions based on carbonate delta238U modeling. However, there have been limited attempts to quantify uncertainty in metal isotope mass balance approaches. Here, we probabilistically evaluate coupled metal isotopes and sedimentary archives to increase constraint. We present iron speciation, metal concentration, delta98Mo and delta238U measurements of Rhuddanian black shales from the Murzuq Basin, Libya. We evaluate these data (and published carbonate delta238U data) with a coupled stochastic mass balance model. Combined statistical analysis of metal isotopes and sedimentary sinks provides uncertainty-bounded constraints on the intensity of Hirnantian-Rhuddanian euxinia. This work extends the duration of anoxia to >3 Myrs - notably longer than well-studied Mesozoic ocean anoxic events.

    View details for DOI 10.1038/s41467-020-15400-y

    View details for PubMedID 32286253

  • On the co-evolution of surface oxygen levels and animals. Geobiology Cole, D. B., Mills, D. B., Erwin, D. H., Sperling, E. A., Porter, S. M., Reinhard, C. T., Planavsky, N. J. 2020


    Few topics in geobiology have been as extensively debated as the role of Earth's oxygenation in controlling when and why animals emerged and diversified. All currently described animals require oxygen for at least a portion of their life cycle. Therefore, the transition to an oxygenated planet was a prerequisite for the emergence of animals. Yet, our understanding of Earth's oxygenation and the environmental requirements of animal habitability and ecological success is currently limited; estimates for the timing of the appearance of environments sufficiently oxygenated to support ecologically stable populations of animals span a wide range, from billions of years to only a few million years before animals appear in the fossil record. In this light, the extent to which oxygen played an important role in controlling when animals appeared remains a topic of debate. When animals originated and when they diversified are separate questions, meaning either one or both of these phenomena could have been decoupled from oxygenation. Here, we present views from across this interpretive spectrum-in a point-counterpoint format-regarding crucial aspects of the potential links between animals and surface oxygen levels. We highlight areas where the standard discourse on this topic requires a change of course and note that several traditional arguments in this "life versus environment" debate are poorly founded. We also identify a clear need for basic research across a range of fields to disentangle the relationships between oxygen availability and emergence and diversification of animal life.

    View details for DOI 10.1111/gbi.12382

    View details for PubMedID 32175670

  • Sources of C30 steroid biomarkers in Neoproterozoic-Cambrian rocks and oils. Nature ecology & evolution Love, G. D., Zumberge, J. A., Cardenas, P., Sperling, E. A., Rohrssen, M., Grosjean, E., Grotzinger, J. P., Summons, R. E. 2019

    View details for DOI 10.1038/s41559-019-1048-2

    View details for PubMedID 31768019

  • New insights on the Orosirian carbon cycle, early Cyanobacteria, and the assembly of Laurentia from the Paleoproterozoic Belcher Group EARTH AND PLANETARY SCIENCE LETTERS Hodgskiss, M. W., Dagnaud, O. J., Frost, J. L., Halverson, G. P., Schmitz, M. D., Swanson-Hysell, N. L., Sperling, E. A. 2019; 520: 141–52
  • Statistical inference and reproducibility in geobiology GEOBIOLOGY Sperling, E. A., Tecklenburg, S., Duncan, L. E. 2019; 17 (3): 261–71

    View details for DOI 10.1111/gbi.12333

    View details for Web of Science ID 000465014600003

  • Oxygen, temperature and the deep-marine stenothermal cradle of Ediacaran evolution. Proceedings. Biological sciences Boag, T. H., Stockey, R. G., Elder, L. E., Hull, P. M., Sperling, E. A. 2018; 285 (1893): 20181724


    Ediacaran fossils document the early evolution of complex megascopic life, contemporaneous with geochemical evidence for widespread marine anoxia. These data suggest early animals experienced frequent hypoxia. Research has thus focused on the concentration of molecular oxygen (O2) required by early animals, while also considering the impacts of climate. One model, the Cold Cradle hypothesis, proposed the Ediacaran biota originated in cold, shallow-water environments owing to increased O2 solubility. First, we demonstrate using principles of gas exchange that temperature does have a critical role in governing the bioavailability of O2-but in cooler water the supply of O2 is actually lower. Second, the fossil record suggests the Ediacara biota initially occur approximately 571 Ma in deep-water facies, before appearing in shelf environments approximately 555 Ma. We propose an ecophysiological underpinning for this pattern. By combining oceanographic data with new respirometry experiments we show that in the shallow mixed layer where seasonal temperatures fluctuate widely, thermal and partial pressure ( pO2) effects are highly synergistic. The result is that temperature change away from species-specific optima impairs tolerance to low pO2. We hypothesize that deep and particularly stenothermal (narrow temperature range) environments in the Ediacaran ocean were a physiological refuge from the synergistic effects of temperature and low pO2.

    View details for DOI 10.1098/rspb.2018.1724

    View details for PubMedID 30963899

    View details for PubMedCentralID PMC6304043

  • Oxygenated Mesoproterozoic lake revealed through magnetic mineralogy. Proceedings of the National Academy of Sciences of the United States of America Slotznick, S. P., Swanson-Hysell, N. L., Sperling, E. A. 2018


    Terrestrial environments have been suggested as an oxic haven for eukaryotic life and diversification during portions of the Proterozoic Eon when the ocean was dominantly anoxic. However, iron speciation and Fe/Al data from the ca. 1.1-billion-year-old Nonesuch Formation, deposited in a large lake and bearing a diverse assemblage of early eukaryotes, are interpreted to indicate persistently anoxic conditions. To shed light on these distinct hypotheses, we analyzed two drill cores spanning the transgression into the lake and its subsequent shallowing. While the proportion of highly reactive to total iron (FeHR/FeT) is consistent through the sediments and typically in the range taken to be equivocal between anoxic and oxic conditions, magnetic experiments and petrographic data reveal that iron exists in three distinct mineral assemblages resulting from an oxycline. In the deepest waters, reductive dissolution of iron oxides records an anoxic environment. However, the remainder of the sedimentary succession has iron oxide assemblages indicative of an oxygenated environment. At intermediate water depths, a mixed-phase facies with hematite and magnetite indicates low oxygen conditions. In the shallowest waters of the lake, nearly every iron oxide has been oxidized to its most oxidized form, hematite. Combining magnetics and textural analyses results in a more nuanced understanding of ambiguous geochemical signals and indicates that for much of its temporal duration, and throughout much of its water column, there was oxygen in the waters of Paleolake Nonesuch.

    View details for PubMedID 30509974

  • Oxygen, temperature and the deep-marine stenothermal cradle of Ediacaran evolution PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES Boag, T. H., Stockey, R. G., Elder, L. E., Hull, P. M., Sperling, E. A. 2018; 285 (1893)
  • Demosponge steroid biomarker 26-methylstigmastane provides evidence for Neoproterozoic animals. Nature ecology & evolution Zumberge, J. A., Love, G. D., Cardenas, P., Sperling, E. A., Gunasekera, S., Rohrssen, M., Grosjean, E., Grotzinger, J. P., Summons, R. E. 2018; 2 (11): 1709–14


    Sterane biomarkers preserved in ancient sedimentary rocks hold promise for tracking the diversification and ecological expansion of eukaryotes. The earliest proposed animal biomarkers from demosponges (Demospongiae) are recorded in a sequence around 100Myr long of Neoproterozoic-Cambrian marine sedimentary strata from the Huqf Supergroup, South Oman Salt Basin. This C30 sterane biomarker, informally known as 24-isopropylcholestane (24-ipc), possesses the same carbon skeleton as sterols found in some modern-day demosponges. However, this evidence is controversial because 24-ipc is not exclusive to demosponges since 24-ipc sterols are found in trace amounts in some pelagophyte algae. Here, we report a new fossil sterane biomarker that co-occurs with 24-ipc in a suite of late Neoproterozoic-Cambrian sedimentary rocks and oils, which possesses a rare hydrocarbon skeleton that is uniquely found within extant demosponge taxa. This sterane is informally designated as 26-methylstigmastane (26-mes), reflecting the very unusual methylation at the terminus of the steroid side chain. It is the first animal-specific sterane marker detected in the geological record that can be unambiguously linked to precursor sterols only reported from extant demosponges. These new findings strongly suggest that demosponges, and hence multicellular animals, were prominent in some late Neoproterozoic marine environments at least extending back to the Cryogenian period.

    View details for PubMedID 30323207

  • Demosponge steroid biomarker 26-methylstigmastane provides evidence for Neoproterozoic animals NATURE ECOLOGY & EVOLUTION Zumberge, J., Love, G. D., Cardenas, P., Sperling, E. A., Gunasekera, S., Rohrssen, M., Grosjean, E., Grotzinger, J. P., Summons, R. E. 2018; 2 (11): 1709–14
  • The Temporal and Environmental Context of Early Animal Evolution: Considering All the Ingredients of an "Explosion". Integrative and comparative biology Sperling, E. A., Stockey, R. G. 2018; 58 (4): 605–22


    Animals originated and evolved during a unique time in Earth history-the Neoproterozoic Era. This paper aims to discuss (1) when landmark events in early animal evolution occurred, and (2) the environmental context of these evolutionary milestones, and how such factors may have affected ecosystems and body plans. With respect to timing, molecular clock studies-utilizing a diversity of methodologies-agree that animal multicellularity had arisen by 800 million years ago (Ma) (Tonian period), the bilaterian body plan by 650 Ma (Cryogenian), and divergences between sister phyla occurred 560-540 Ma (late Ediacaran). Most purported Tonian and Cryogenian animal body fossils are unlikely to be correctly identified, but independent support for the presence of pre-Ediacaran animals is recorded by organic geochemical biomarkers produced by demosponges. This view of animal origins contrasts with data from the fossil record, and the taphonomic question of why animals were not preserved (if present) remains unresolved. Neoproterozoic environments demanding small, thin, body plans, and lower abundance/rarity in populations may have played a role. Considering environmental conditions, geochemical data suggest that animals evolved in a relatively low-oxygen ocean. Here, we present new analyses of sedimentary total organic carbon contents in shales suggesting that the Neoproterozoic ocean may also have had lower primary productivity-or at least lower quantities of organic carbon reaching the seafloor-compared with the Phanerozoic. Indeed, recent modeling efforts suggest that low primary productivity is an expected corollary of a low-O2 world. Combined with an inability to inhabit productive regions in a low-O2 ocean, earliest animal communities would likely have been more food limited than generally appreciated, impacting both ecosystem structure and organismal behavior. In light of this, we propose the "fire triangle" metaphor for environmental influences on early animal evolution. Moving toward consideration of all environmental aspects of the Cambrian radiation (fuel, heat, and oxidant) will ultimately lead to a more holistic view of the event.

    View details for PubMedID 30295813

  • On the edge of exceptional preservation: insights into the role of redox state in Burgess Shale-type taphonomic windows from the Mural Formation, Alberta, Canada. Emerging topics in life sciences Sperling, E. A., Balthasar, U., Skovsted, C. B. 2018; 2 (2): 311-323


    Animals originated in the Neoproterozoic and 'exploded' into the fossil record in the Cambrian. The Cambrian also represents a high point in the animal fossil record for the preservation of soft tissues that are normally degraded. Specifically, fossils from Burgess Shale-type (BST) preservational windows give paleontologists an unparalleled view into early animal evolution. Why this time interval hosts such exceptional preservation, and why this preservational window declines in the early Paleozoic, have been long-standing questions. Anoxic conditions have been hypothesized to play a role in BST preservation, but recent geochemical investigations of these deposits have reached contradictory results with respect to the redox state of overlying bottom waters. Here, we report a multi-proxy geochemical study of the Lower Cambrian Mural Formation, Alberta, Canada. At the type section, the Mural Formation preserves rare recalcitrant organic tissues in shales that were deposited near storm wave base (a Tier 3 deposit; the worst level of soft-tissue preservation). The geochemical signature of this section shows little to no evidence of anoxic conditions, in contrast with published multi-proxy studies of more celebrated Tier 1 and 2 deposits. These data help confirm that 'decay-limited' BST biotas were deposited in more oxygenated conditions, and support a role for anoxic conditions in BST preservation. Finally, we discuss the role of iron reduction in BST preservation, including the formation of iron-rich clays and inducement of sealing seafloor carbonate cements. As oceans and sediment columns became more oxygenated and more sulfidic through the early Paleozoic, these geochemical changes may have helped close the BST taphonomic window.

    View details for DOI 10.1042/ETLS20170163

    View details for PubMedID 32412614

  • Temperature-dependent hypoxia explains biogeography and severity of end-Permian marine mass extinction. Science (New York, N.Y.) Penn, J. L., Deutsch, C., Payne, J. L., Sperling, E. A. 2018; 362 (6419)


    Rapid climate change at the end of the Permian Period (~252 million years ago) is the hypothesized trigger for the largest mass extinction in Earth's history. We present model simulations of the Permian/Triassic climate transition that reproduce the ocean warming and oxygen (O2) loss indicated by the geologic record. The effect of these changes on animal survival is evaluated using the Metabolic Index (Phi), a measure of scope for aerobic activity governed by organismal traits sampled in diverse modern species. Modeled loss of aerobic habitat predicts lower extinction intensity in the tropics, a pattern confirmed with a spatially explicit analysis of the marine fossil record. The combined physiological stresses of ocean warming and O2 loss can account for more than half the magnitude of the "Great Dying."

    View details for PubMedID 30523082

  • Oxygen, facies, and secular controls on the appearance of Cryogenian and Ediacaran body and trace fossils in the Mackenzie Mountains of northwestern Canada GEOLOGICAL SOCIETY OF AMERICA BULLETIN Sperling, E. A., Carbone, C., Strauss, J. V., Johnston, D. T., Narbonne, G. M., Macdonald, F. A. 2016; 128 (3-4): 558-575

    View details for DOI 10.1130/B31329.1

    View details for Web of Science ID 000370973000012

  • Biotic replacement and mass extinction of the Ediacara biota. Proceedings. Biological sciences / The Royal Society Darroch, S. A., Sperling, E. A., Boag, T. H., Racicot, R. A., Mason, S. J., Morgan, A. S., Tweedt, S., Myrow, P., Johnston, D. T., Erwin, D. H., Laflamme, M. 2015; 282 (1814)


    The latest Neoproterozoic extinction of the Ediacara biota has been variously attributed to catastrophic removal by perturbations to global geochemical cycles, 'biotic replacement' by Cambrian-type ecosystem engineers, and a taphonomic artefact. We perform the first critical test of the 'biotic replacement' hypothesis using combined palaeoecological and geochemical data collected from the youngest Ediacaran strata in southern Namibia. We find that, even after accounting for a variety of potential sampling and taphonomic biases, the Ediacaran assemblage preserved at Farm Swartpunt has significantly lower genus richness than older assemblages. Geochemical and sedimentological analyses confirm an oxygenated and non-restricted palaeoenvironment for fossil-bearing sediments, thus suggesting that oxygen stress and/or hypersalinity are unlikely to be responsible for the low diversity of communities preserved at Swartpunt. These combined analyses suggest depauperate communities characterized the latest Ediacaran and provide the first quantitative support for the biotic replacement model for the end of the Ediacara biota. Although more sites (especially those recording different palaeoenvironments) are undoubtedly needed, this study provides the first quantitative palaeoecological evidence to suggest that evolutionary innovation, ecosystem engineering and biological interactions may have ultimately caused the first mass extinction of complex life.

    View details for DOI 10.1098/rspb.2015.1003

    View details for PubMedID 26336166

    View details for PubMedCentralID PMC4571692

  • Statistical analysis of iron geochemical data suggests limited late Proterozoic oxygenation NATURE Sperling, E. A., Wolock, C. J., Morgan, A. S., Gill, B. C., Kunzmann, M., Halverson, G. P., Macdonald, F. A., Knoll, A. H., Johnston, D. T. 2015; 523 (7561): 451-454
  • The Ecological Physiology of Earth's Second Oxygen Revolution ANNUAL REVIEW OF ECOLOGY, EVOLUTION, AND SYSTEMATICS, VOL 46 Sperling, E. A., Knoll, A. H., Girguis, P. R. 2015; 46: 215-235
  • A Permian-Triassic boundary section at Quinn River Crossing, northwestern Nevada, and implications for the cause of the Early Triassic chert gap on the western Pangean margin GEOLOGICAL SOCIETY OF AMERICA BULLETIN Sperling, E. A., Ingle, J. C. 2006; 118 (5-6): 733-746

    View details for DOI 10.1130/B25803.1

    View details for Web of Science ID 000237310200015