Earth and Planetary Science Letters 361 (2013) 429–435 Contents lists available at SciVerse ScienceDirect Earth and Planetary Science Letters journal homepage: www.elsevier.com/locate/epsl Constraints on Early Triassic carbon cycle dynamics from paired organic and inorganic carbon isotope records K.M. Meyer a,n, M. Yu b, D. Lehrmann c, B. van de Schootbrugge d, J.L. Payne a a Department of Geological & Environmental Sciences, Stanford University, 450 Serra Mall, Building 320, Stanford, CA 94305, USA College of Resource and Environment Engineering, Guizhou University, Caijiaguan, Guiyang 550003, Guizhou Province, PR China c Department of Geoscience, Trinity University, One Trinity Place, San Antonio, TX 78212, USA d Institute of Geosciences, Johann Wolfgang Goethe University Frankfurt, Altenhöferallee 1, Frankfurt am Main D-60438, Germany b a r t i c l e i n f o abstract Article history: Received 5 June 2012 Received in revised form 22 October 2012 Accepted 26 October 2012 Editor: J. Lynch-Stieglitz Available online 5 December 2012 Large d13C excursions, anomalous carbonate precipitates, low diversity assemblages of small fossils, and evidence for marine euxinia in uppermost Permian and Lower Triassic strata bear more similarity to Neoproterozoic carbonates than to the remainders of the Permian and Triassic systems. Middle Triassic diversification of marine ecosystems coincided with the waning of anoxia and stabilization of the global carbon cycle, suggesting that environment-ecosystem linkages were important to biological recovery. However, the Earth system behavior responsible for these large d13C excursions remains poorly constrained. Here we present a continuous Early Triassic d13Corg record from south China and use it to test the extent to which Early Triassic excursions in d13Ccarb record changes in the d13C of marine dissolved inorganic carbon (DIC). Regression analysis demonstrates a significant positive correlation between d13Corg and d13Ccarb across multiple sections that span a paleoenvironmental gradient. Such a correlation is incompatible with diagenetic alteration because no likely mechanism will alter both d13Corg and d13Ccarb records in parallel and therefore strongly indicates a primary depositional origin. A simple explanation for this correlation is that a substantial portion of the preserved Corg was derived from the contemporaneous DIC pool, implying that the observed excursions reflect variation in the d13C of the exogenic carbon reservoir (ocean, atmosphere, biomass). These findings support existing evidence that large d13C excursions are primary and therefore strengthen the case that large-scale changes to the carbon cycle were mechanistically linked to the low diversity and small size of Early Triassic fossils. Associated sedimentary and biogeochemical phenomena further suggest that similar associations in Neoproterozoic and Cambrian strata may reflect the same underlying controls. & 2012 Elsevier B.V. All rights reserved. Keywords: Permian–Triassic boundary extinction stable carbon isotopes China carbon cycle 1. Introduction Some of the largest d13C excursions of the Phanerozoic are recorded in Lower Triassic limestones, deposited during the aftermath of the end-Permian mass extinction (Payne et al., 2004). These large amplitude ( 48%) positive and negative shifts are similar in magnitude and timescale to the enigmatic Neoproterozoic and Cambrian carbon isotope perturbations (Halverson et al., 2005; Maloof et al., 2010). The similarities extend beyond chemical signatures to characteristic carbonate sediment fabrics, exceptionally low animal abundance, low taxonomic diversity, small organism size, and evidence for widespread ocean anoxia and euxinia (Grotzinger and Knoll, 1995; Maloof et al., 2010; Meyer et al., 2011; Payne et al., 2004). These observations suggest n Corresponding author. E-mail address: meyerk@stanford.edu (K.M. Meyer). 0012-821X/$ - see front matter & 2012 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.epsl.2012.10.035 intimate relationships between carbon cycle behavior, biodiversity, and ecosystem function for both Neoproterozoic–Cambrian and Early Triassic intervals (Maloof et al., 2005, 2010). However, the causes of very large and rapid ( o1 My) carbon isotope excursions in marine limestones remain poorly understood and several of the proposed explanations are related to diagenesis or spatial variation in carbonate mineralogy rather than disturbances of the global carbon cycle (Derry, 2010; Swart and Eberli, 2005). Consequently, constraining the causes of large variations in the d13C of inorganic carbon in carbonate rocks is central to understanding the mechanistic links, if any, between the Early Triassic sedimentary, paleobiological, and biogeochemical records. Early Triassic excursions in the d13C of carbonate rocks 13 (d Ccarb) occur across the Tethys (Atudorei, 1999; Horacek et al., 2007; Payne et al., 2004; Richoz, 2004) and in records from Panthalassa (Horacek et al., 2009) and have therefore been widely interpreted to result from variation in the d13C of Early Triassic 430 K.M. Meyer et al. / Earth and Planetary Science Letters 361 (2013) 429–435 107° E Guiyang 26° N 30°N Yangtze Block Guanling Great Bank of Guizhou N Nanpanjiang Basin 110°E tze fo lat rm Zhenfeng P Gu g n Ya izh Luodian ou a Gu i x ng S Nanpanjiang Basin N S M. Triassic L.Triassic U. Permian Bianyang Guandao Dawen 250 m Shelly limestone Tubiphytes Reef 1 km Oolitic limestone Peritidal cyclic limestone and dolomite Pelagic limestone and allodapic breccias Shale Stratigraphic section Thrombolite-bearing cyclic limestone Fig. 1. (A) Paleogeographic map of the Early Triassic GBG (Lehrmann et al., 1998). Inset: The stippled pattern indicates the Nanpanjiang Basin and the brick pattern represents the Yangtze Block. (B) Schematic cross section of the GBG, modified after Lehrmann et al. (1998). The vertical bars illustrate the locations of the stratigraphic sections within the GBG. dissolved inorganic carbon (DIC) in the oceans. Carbon isotope stabilization early in the Anisian (Middle Triassic) has been interpreted to reflect either the end of environmental forcing of carbon cycle perturbations or a stabilization of the carbon cycle caused by the recovery of global ecosystems (Payne et al., 2004). Like their Early Triassic counterparts, many Precambrian– Cambrian isotope excursions can be correlated over vast distances but the underlying causes remain widely debated. They have been variously interpreted to reflect perturbations of the global carbon cycle, unusual configurations or sizes of carbon reservoirs in the oceans (Grotzinger et al., 2011; Rothman et al., 2003; Tziperman et al., 2011), or diagenetic overprints (Derry, 2010; Knauth and Kennedy, 2009). Global processes, such as sea level change, impact the distribution of carbonate mineralogies and depositional environments and are of particular relevance to the Early Triassic because the d13Ccarb record observed on widely separated carbonate platforms could show similar trends in the absence of a change in the d13C of marine DIC (Swart and Eberli, 2005). Similarly, global changes in the relative importance of diagenetic processes such as methanogenesis could also cause an observed shift in d13Ccarb without a similar change in the d13C of the DIC reservoir (Hayes and Waldbauer, 2006). Thus, the observation of globally correlated isotope excursions does not unequivocally establish that the d13Ccarb variations record secular changes in the d13C of the marine DIC reservoir. The carbon isotope composition of organic matter (d13Corg) has the potential to better constrain the causes of the large Early Triassic C isotope excursions. Most Phanerozoic paired d13Ccarb and d13Corg records display the consistent coupling that is expected when marine organic matter is produced from carbon in isotopic equilibrium with contemporaneous dissolved inorganic carbon (DIC), especially when variation in d13C of DIC is greater than any variability in e (fractionation between d13CDIC and d13Corg during carbon fixation). Any alteration of the d13Corg record would result from processes different from those known to diagenetically alter d13Ccarb (Knoll et al., 1986). For example, thermal maturation of organic matter decreases the total organic carbon (TOC) composition of rocks and tends to shift residual TOC to more enriched d13C values (Hayes et al., 1999). This process would impact the d13Corg values but not the trends (Des Marais et al., 1992). Migration of hydrocarbons or contamination by detrital organic Corg from rock weathering could also affect d13Corg K.M. Meyer et al. / Earth and Planetary Science Letters 361 (2013) 429–435 values. Again, these processes would not impact the d13Ccarb record, and could be evaluated by examining d13Corg values by facies or in comparison to TOC. Therefore, paired d13Ccarb and d13Corg records can reveal the likelihood that perturbations in either one are primary. If paired d13Ccarb and d13Corg measurements are correlated, the record is likely driven by changes in the exogenic carbon cycle. If the isotope measurements are not correlated, the perturbations may be related to diagenetic controls (Derry, 2010; Knauth and Kennedy, 2009) or non-steady state carbon cycle behavior and the buildup and oxidation of a large organic carbon reservoir (Rothman et al., 2003; SwansonHysell et al., 2010). If the Early Triassic records are correlated and the d13Ccarb excursions represent changes in the contemporaneous carbon cycle, then a steady state interpretation of C cycle dynamics requires extreme shifts in the ratio of organic to inorganic carbon burial (Maloof et al., 2010; Payne and Kump, 2007), a substantial change in riverine d13Ccarb, or a change in the fractionation between d13Ccarb and d13Corg. Here we use a paired carbon isotope approach to evaluate the extent to which the observed perturbations reflect changes in the Early Triassic ocean chemistry and to gain insight into the nature of the carbon cycle during biotic recovery from mass extinction. 2. Field setting and methods We measured the d13C composition of limestone and associated organic matter from hand samples from the Great Bank of Guizhou (GBG), an exceptionally exposed carbonate platform in the Nanpanjiang Basin of south China (Fig. 1). A syncline dissects the GBG, providing cross-sectional exposure of thick Upper Permian through Middle Triassic stratigraphic sections along an inferred paleobathymetric gradient from the shallow-water (intertidal to tens of meters) platform interior to deep-water (hundreds of meters) basin margin environments (Lehrmann et al., 1998). Separate sections are correlated based upon field stratigraphic relations and conodont and foraminiferan biostratigraphy (Lehrmann et al., 1998; Meyer et al., 2011; Payne et al., 2004). Geochemical and stable isotope analyses were performed in the Stanford University Stable Isotope Biogeochemistry Laboratories. Limestone samples were powdered using a 0.8 mm dental drill bit. Micrites were selectively sampled, avoiding calcite veins and fossils. Dolomitized intervals of Laolaicao and Dajiang were also sampled, but at a lower density, for completeness. Methods for d13Ccarb analysis are fully described in Meyer et al. (2011). For d13Corg analysis, hand samples were powdered with a masonry bit, with special care taken to avoid weathering fronts and veins. 1–3 g of dried, powdered limestone was dissolved in 3 N hydrochloric acid until reaction was complete to remove carbonate minerals, typically less than 1 h. After centrifugation, the remaining supernatant was removed and each sample was rinsed 3 times with 18.2 MO water. d13Corg values of the dried, powdered residues were then determined using a Costech elemental analyzer coupled to a Thermo Finnegan Deltaplus XL isotope ratio mass spectrometer. Isotopic composition is reported in standard delta notation relative to the VPDB standard. Analytical precision was typically better than 0.2% based on replicate measurements of an internal laboratory standard. 3. Results The Lower Triassic d13Corg records from Dawen, Guandao, and Bianyang presented in Fig. 2 display a magnitude of negative and positive shifts of up to 6–7%, excluding outliers. This pattern 431 follows that of the d13Ccarb record at these sections and is consistent with the d13Ccarb records across the GBG. The d13Corg data display much greater sample-to-sample scatter than the d13Ccarb values and there is no discernable d13Corg gradient across the platform (Fig. 5a). All three sections display shifts in the d13Corg record that correspond to shifts in their respective d13Ccarb records, though the amplitude of d13Corg variation is damped. As a result, the D13C (d13Ccarb–d13Corg) also exhibits shifts parallel to those seen in d13Ccarb (Fig. 2). A positive correlation between organic and inorganic carbon isotopes is apparent in scatterplots of d13Corg versus d13Ccarb for each section (Fig. 3). This correlation is shown to be statistically robust through generalized least squares regression (Table 1), which shows that the slopes of d13Ccarb and d13Corg are statistically different from both zero and one, ranging between 0.4 and 0.6. 4. Discussion The d13Corg data are consistent with several other lines of evidence indicating that the large d13Ccarb excursions are primary, including a lack of significant correlation between d13Ccarb and d18O or trace metals (as shown in Meyer et al. (2011)). The analysis presented above indicates that the major excursions in the d13Ccarb record were not significantly altered by meteoric diagenesis, organic matter remineralization, or changes in the mixing ratio of sediment sources and minerals across the GBG. There is also no correlation between TOC and d13Corg, suggesting that there is no strong diagenetic control of the Corg record (Fig. 4; see also Johnston et al., 2012). Finally, organic petrographic observations show that the organic matter in GBG limestones is primarily amorphous organic matter with a likely bacterial or marine phytoplanktonic origin. There is no evidence for substantial contamination by modern organic matter (Fig. S1). The positive relationship between d13Corg and d13Ccarb arising from coeval positive and negative shifts, the global nature of the d13C carb shifts, and geochemical tests of diagenesis suggest that the inorganic and organic carbon records reflect variation in the isotopic composition of contemporaneous DIC. The regression analysis further supports the coupled nature of these measurements but do not offer a mechanism or physical explanation for the nature of the coupling. Although the d13Corg and d13Ccarb records are correlated, the slope of the relationship is significantly less than unity (Table 1), which is the expected value if organic carbon is formed from contemporaneous DIC and the fractionation associated with photosynthesis is relatively invariant. The imperfect correlation between d13Corg and d13Ccarb could be influenced by contamination, by a large DOC pool and consequent time-averaging of the d13Corg signal, or by the mixing of Corg sources. Here we discuss the relative support for these various possibilities. First, GBG limestones are low in TOC, and some authors have suggested that low TOC samples may be prone to contamination by modern organic matter. However, we find no statistically significant correlation between %TOC and d13Corg (Fig. 4; adjusted R^2¼0.003, p ¼0.21) or between lithofacies and d13Corg (Fig. S2). In addition, organic petrographic analyses of selected GBG samples show no evidence for significant modern material in the sample residues (Fig. S1). These lines of evidence suggest the correlation less than unity in the paired d13Corg and d13Ccarb data is not likely due to contamination by modern organic matter. Variation in d13Corg could alternatively be damped relative to changes in d13Ccarb if there is a large DOC reservoir due to timeaveraging of the d13Corg signal. Rothman et al. (2003) have described a mechanism that can explain decoupling of d13Ccarb and d13Corg through buildup and oxidation of a large DOC 432 K.M. Meyer et al. / Earth and Planetary Science Letters 361 (2013) 429–435 Fig. 2. Organic and inorganic carbon isotopic data from three stratigraphic sections across the GBG, modified after Meyer et al. (2011). The time scale is based on conodont biostratigraphy from Guandao (Payne et al., 2004), and correlations are suggested based upon lithostratigraphic relationships (red) and carbon isotope trends (blue). Conodont first occurrences: a. Neogondolella changxingensis, b. Hindeodus parvus, c. Neospathodus dieneri, d. Ns. waageni, e. Ns. Bansoni. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.) Fig. 3. Relationship between d13Ccarb and d13Corg for Dawen, Guandao, and Bianyang limestones. The coefficients of determination (R2) for linear regressions through these data are 0.43, 0.30, and 0.53, respectively. Full regression statistics are provided in Table 1. Table 1 Generalized least squares regression shows that the slopes of d13Ccarb and d18Corg are statistically different from both 0 and 1 and range between 0.4 and 0.6. Section Intercept (e) Standard error Slope Standard error p Bianyang Dawen Guandao Upper Guandao 28.6 28.2 27.8 28.3 0.22 0.15 0.30 0.68 0.52 0.41 0.48 0.69 0.09 0.05 0.10 0.29 o0.0001 o0.0001 o0.0001 0.04 reservoir. This non-steady state approach to C cycle dynamics has been used to interpret the Neoproterozoic Shuram excursion, where a large carbon isotope excursion observed in d13Ccarb is absent in d13Corg (Fike et al., 2006). Oxidation of a very large marine DOC pool relative to DIC would cause a negative isotope excursion in d13Ccarb without changing d13Corg. Based on visual inspection of values calculated using Rothman et al.’s (2003) equations, we find that the DOC:DIC ratio would have to be at least 4 to fit the slope we observe. In addition, the relationship between d13Ccarb and e (difference between d13Ccarb and d13Corg) is nearly indistinguishable from the relationship shown by Rothman et al. (2003) for 731–590 Ma. Although the extreme scenario for the Shuram has not been proposed for the Permian– Triassic, non-linear carbon cycle dynamics and the oxidation of marine organic matter have been implicated in the negative carbon isotope excursion at the P–T boundary (Rothman, 2010). Others have suggested that a vigorous biological pump (Meyer et al., 2011) and DOC oxidation coupled to sulfate reduction may have impacted C and S cycling during this interval (Luo et al., K.M. Meyer et al. / Earth and Planetary Science Letters 361 (2013) 429–435 2010). The key question that remains for this scenario is the uncertain timescale over which DOC buildup would have been possible. 0.00 0.05 0.10 0.15 TOC 0.20 0.25 0.30 Alternatively, if the Corg record was originally derived from the DIC pool, then the observed decoupling (slopes less than one in Table 1) between the two isotope records could reflect either covariation of photosynthetic fractionation with the d13C of DIC or the mixing of two pools of organic carbon, one of which was tracking the d13C of DIC with a constant offset due to photosynthetic fractionation and one of which was invariant or varying independently of the d13C of DIC. The former possibility is consistent with the fact that the photosynthetic fractionation varies over time due to changes in pCO2, growth rate, cell geometry, carbon concentrating mechanism, and other factors (reviewed by Hayes et al. (1999)). Given the many degrees of freedom, it is difficult to either rule out this possibility or to demonstrate clearly that it is the most likely explanation for the observed isotopic variations. The latter possibility can be quantified and evaluated more easily, as discussed below. Variation in the d13Corg signal may be damped relative to variation in the d13Ccarb signal by the addition of another pool of organic carbon with a temporally fixed d13C composition, such as recycled ancient organic carbon (weathered TOC), marine DOC (when this reservoir is large relative to DIC), or modern terrestrial organic matter. Recently, Johnston et al. (2012) proposed the incorporation of recycled Corg into the d13Corg signal as an explanation for intervals where the Neoproterozoic d13Corg and d13Corg records become uncorrelated (because the d13Corg record shows little or no stratigraphically coherent variation). To quantify the conditions under which this scenario would account for our data, we performed a second regression analysis to estimate the isotope composition and proportional contribution of a second carbon pool to the measured d13Corg record. 0.35 -21 -23 -25 -27 -29 -31 -33 -35 Dajiang Daw en Cordev. Late Triassic Carnian Fig. 4. Relationship between d18Corg and TOC for Dawen, Guandao, and Bianyang sections (R2 ¼ 0.09). 236 433 Laolaicao Daw en Guandao Bianyang Guandao Bianyang Ladinian 238 240 Long. 242 Middle Triassic 244 246 Pelsonian Anisian Illyrian A./B. 250 Early Triassic Spathian 248 Olenekian Age (millions of years before present) Fass. Smith. Ind. 252 Late Permian Dien. Gries. Late Changsingian -4 -2 0 C 2 4 6 (‰ VPDB) 8 -4 -2 0 C 2 4 6 (‰ VPDB) 8 -36 -34 -32 -30 -28 -26 -24 C (‰ VPDB) 0 5 10 15 20 Observed diversity persample 0 1 20 30 40 50 60 Skeletal abundance as vol % Fig. 5. Summary of Triassic C isotope and paleobiological trends. (A) The Early and Middle Triassic d13C curve from all GBG sections. (B and C) The Early and Middle Triassic d13Corg and d13Ccarb gradients across the GBG. Plotted d13C values are from correlated samples across the slope (filled symbols) and platform interior sections (open symbols) as in Meyer et al. (2011). The d13Ccarb gradient is about 4% in the Early Triassic. It collapses to approximately zero in the Middle Triassic interval of accelerated biotic recovery and carbon isotope stabilization. In contrast, there is no coherent gradient in d13Corg observed across the GBG during the Early Triassic. (D and E) Box plots of diversity and fossil abundance observed on the GBG from the Late Permian into the Late Triassic. Data from Payne et al. (2006). 434 K.M. Meyer et al. / Earth and Planetary Science Letters 361 (2013) 429–435 Assuming a constant 30% fractionation during photosynthesis, the equation to describe the proportional contribution of two organic carbon pools to the measured d13Corg values is: d13 Corg ¼ ðd13 Ccarb 30Þ f contemp þ ð1f contemp Þ d13 Cother where fcontemp represents the fraction of Corg derived from the contemporaneous marine Corg pool and d13Cother represents the isotope composition of carbon derived from a second pool (weathered organic carbon, terrestrial organic carbon, modern contamination, etc.). Based on this model for the paired d13Corg and d13Ccarb data across the GBG, regression analysis suggests that 45% of the Corg measured was derived from the contemporaneous DIC pool (for Bianyang fcontemp ¼0.52, std error¼ 0.10; for Dawen, fcontemp ¼0.41, std error¼0.17; for Guandao, fcontemp ¼0.48, std error¼0.10). This analysis implies that over half of the Corg must have been from another source with a stochastically constant composition. Such alternative sources of Corg include weathered carbon, terrestrial inputs from a variety of sources, and a large DOC pool with relatively invariant d13C. The predicted d13C composition of this non-DIC derived pool from regression modeling is 27%. Unfortunately, the isotope composition of the second carbon pool does not allow us to distinguish between these sources. Regardless of the source of this other organic carbon pool, however, the significant correlation between the d13Corg and d13Ccarb records supports a causal connection between carbon isotope stabilization and enhanced biotic recovery in Middle Triassic time; carbon isotope stabilization is associated with increases in fossil diversity and abundance across the platform (Fig. 5). At present, we favor the scenario in which the bulk organic carbon isotope record reflects the mixing of carbon from two different sources. The extremely low TOC of our samples means that even a small amount of organic carbon from a source other than contemporaneous marine organisms could have a substantial impact on the bulk d13Corg record. The larger sample-tosample variation in d13Corg versus d13Ccarb is also consistent with variation among samples in the proportion of contemporaneous versus other organic carbon. We cannot rule out the possibility that temporal variation in the photosynthetic fractionation of carbon isotopes also contributed to the reduced variation in d13Corg, but under this scenario our data require that the factors influencing the photosynthetic fractionation covaried with d13Ccarb with a slope of approximately one half. 5. Conclusions and implications Here we present the first paired d13Ccarb and d13Corg record for the latest Permian and Early Triassic. Large positive and negative excursions occur in both records, but the magnitude of the shift is slightly damped in d13Corg. Regression modeling demonstrates a significant correlation between d13Corg and d13Ccarb, requiring that a significant fraction of the preserved Corg formed from the contemporaneous DIC pool. This result therefore implies that the observed excursions reflect variation in the d13C of the exogenic carbon reservoir. In addition, this evidence further implies that Early and Middle Triassic changes in carbon cycle behavior, carbonate deposition, and marine ecosystem recovery share underlying controls. Parallels between these findings and recent results from Neoproterozoic strata also suggest that a damped response in d13Corg relative to d13Ccarb may be typical of low TOC rocks across high-amplitude carbon isotope excursions of all ages. Acknowledgments The authors thank E. Schaal, X. Li, and H. Fu for assistance in the field. P. Blisnuik provided laboratory assistance. Meyer acknowledges support from the Agouron Institute. This work was supported by NSF grant EAR-0807377-007, ACS PRF grant 45329-G8, and NGS grant 8102-06. Appendix A. Supplementary materials Supplementary data associated with this article can be found in the online version at http://dx.doi.org/10.1016/j.epsl.2012.10.035. References Atudorei, V., 1999. Constraints on the Upper Permian to Upper Triassic Marine Carbon Isotope Curve: Case Studies from the Tethys. Ph.D., University of Lausanne. Derry, L.A., 2010. A burial diagenesis origin for the Ediacaran Shuram–Wonoka carbon isotope anomaly. Earth Planet. Sci. Lett. 294, 152–162. Des Marais, D.J., Strauss, H., Summons, R.E., Hayes, J.M., 1992. Carbon isotope evidence for the stepwise oxidation of the Proterozoic environment. Nature 359, 605–609. Fike, D.A., Grotzinger, J.P., Pratt, L.M., Summons, R.E., 2006. Oxidation of the Ediacaran Ocean. Nature 444, 744–747. Grotzinger, J.P., Fike, D.A., Fischer, W.W., 2011. Enigmatic origin of the largestknown carbon isotope excursion in Earth’s history. Nat. Geosci. 4, 285–292. Grotzinger, J.P., Knoll, A.H., 1995. Anomalous carbonate precipitates: is the Precambrian the key to the Permian? Palaios 10, 578–596. Halverson, G.P., Hoffman, P.F., Schrag, D.P., Maloof, A.C., Rice, A.H.N., 2005. Toward a Neoproterozoic composite carbon-isotope record. Geol. Soc. Am. Bull. 117, 1181–1207. Hayes, J.M., Strauss, H., Kaufman, A.J., 1999. The abundance of 13C in marine organic matter and isotopic fractionation in the global biogoechemical cycle of carbon during the past 800 Ma. Chem. Geol. 161, 103–125. Hayes, J.M., Waldbauer, J.R., 2006. The carbon cycle and associated redox processes through time. Philos. Trans. R. Soc. B-Biol. Sci. 361, 931–950. Horacek, M., Brandner, R., Abart, R., 2007. Carbon isotope record of the P/T boundary and the Lower Triassic in the Southern Alps: evidence for rapid changes in storage of organic carbon. Palaeogeogr. Palaeoclimatol. Palaeoecol. 252, 347–354. Horacek, M., Koike, T., Richoz, S., 2009. Lower Triassic d13C isotope curve from shallow-marine carbonates in Japan, Panthalassa realm: confirmation of the Tethys d13C curve. J. Asian Earth Sci. 36, 481–490. Johnston, D.T., Macdonald, F.A., Gill, B.C., Hoffman, P.F., Schrag, D.P., 2012. Uncovering the Neoproterozoic carbon cycle. Nature 483, 320-U110. Knauth, L.P., Kennedy, M.J., 2009. The late Precambrian greening of the Earth. Nature 460, 728–732. Knoll, A.H., Hayes, J.M., Kaufman, A.J., Swett, K., Lambert, I.B., 1986. Secular variation in carbon isotope ratios from Upper Proterozoic successions of Svalbard and East Greenland. Nature 321, 832–838. Lehrmann, D.J., Wei, J.Y., Enos, P., 1998. Controls on facies architecture of a large Triassic carbonate platform: the Great Bank of Guizhou, Nanpanjiang Basin, South China. J. Sediment. Res. 68, 311–326. Luo, G.M., Kump, L.R., Wang, Y.B., Tong, J.N., Arthur, M.A., Yang, H., Huang, J.H., Yin, H.F., Xie, S.C., 2010. Isotopic evidence for an anomalously low oceanic sulfate concentration following end-Permian mass extinction. Earth Planet. Sci. Lett. 300, 101–111. Maloof, A.C., Porter, S.M., Moore, J.L., Dudas, F.O., Bowring, S.A., Higgins, J.A., Fike, D.A., Eddy, M.P., 2010. The earliest Cambrian record of animals and ocean geochemical change. Geol. Soc. Am. Bull. 122, 1731–1774. Maloof, A.C., Schrag, D.P., Crowley, J.L., Bowring, S.A., 2005. An expanded record of Early Cambrian carbon cycling from the Anti-Atlas Margin, Morocco. Can. J. Earth Sci. 42, 2195–2216. Meyer, K.M., Yu, M., Jost, A.B., Kelley, B.M., Payne, J.L., 2011. d13C evidence that high primary productivity delayed recovery from end-Permian mass extinction. Earth Planet. Sci. Lett. 302, 378–384. Payne, J.L., Kump, L.R., 2007. Evidence for recurrent Early Triassic massive volcanism from quantitative interpretation of carbon isotope fluctuations. Earth Planet. Sci. Lett. 256, 264–277. Payne, J.L., Lehrmann, D.J., Wei, J.Y., Orchard, M.J., Schrag, D.P., Knoll, A.H., 2004. Large perturbations of the carbon cycle during recovery from the end-Permian extinction. Science 305, 506–509. Payne, J.L., Lehrmann, D.J., Wei, J.Y., Knoll, A.H., 2006. The pattern and timing of biotic recovery from the end-Permian extinction on the Great Bank of Guizhou, Guizhou province, China. Palaios 21, 63–85. Richoz, S., 2004. Stratigraphy and Carbon isotope study of the Upper Permian and Lower Triassic in Some Localities of the Neotethyan Realm (Turkey, Oman, Iran), University of Lausanne. Rothman, D.H., 2010. Singular blow-up in the end-Permian carbon cycle, 2010 Fall Meeting. AGU, San Francisco, CA. K.M. Meyer et al. / Earth and Planetary Science Letters 361 (2013) 429–435 Rothman, D.H., Hayes, J.M., Summons, R.E., 2003. Dynamics of the Neoproterozoic carbon cycle. Proc. Natl. Acad. Sci. USA 100, 8124–8129. Swanson-Hysell, N.L., Rose, C.V., Calmet, C.C., Halverson, G.P., Hurtgen, M.T., Maloof, A.C., 2010. Cryogenian glaciation and the onset of carbon-isotope decoupling. Science 328, 608–611. Swart, P.K., Eberli, G., 2005. The nature of the d13C of periplatform sediments: implications for stratigraphy and the global carbon cycle. Sediment. Geol. 175, 115–129. 435 Tziperman, E., Halevy, I., Johnston, D.T., Knoll, A.H., Schrag, D.P., 2011. Biologically induced initiation of Neoproterozoic snowball-Earth events. Proc. Natl. Acad. Sci. USA 108, 15091–15096.