The interaction of recovery and environmental conditions: An analysis of the outer shelf edge of western North America during the early Triassic

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Highlights

  • The outer shelf to slope facies of the Union Wash Formation contain intervals of biotic recovery and environmental stress.
  • Levels of recovery within the Union Wash Formation decrease over time and reflect the influence of environmental stress on recovery.
  • Areas with persistent environmental stress demonstrate long, gradual recoveries, or multiple terminated recoveries.
  • Studies that note rapid recovery within single localities reflect deposition within the habitable zone and do not represent overall recovery.
  • Regional studies provide the most accurate measure of recovery since they take into account the influence environmental variations over broad areas.

Abstract

Biotic recovery from the Permian-Triassic mass extinction was complex and uneven, with prior studies typically revealing a long, drawn-out recovery that was either delayed for some interval, or was subdued and stepwise in nature. Examples of rapid recovery at the outcrop scale cloud this narrative and point to the importance of environmental stresses in determining the timing and shape of recovery. The Union Wash Formation at the Darwin locality (east-central California) contains at least 3 recovery intervals that terminate with the onset of deleterious conditions and therefore offers a means to examine the relationship between environmental stress and recovery. The first recovery is manifested as bioturbated (ii = 5–6), shallow marine micritic limestones that contain small diameter (2–4 mm) Thalassinoides burrows that form complex networks. This interval is overlain by 650.5 m of laminated mudstone that signal deleterious environmental conditions. The second recovery interval begins directly above seafloor precipitate-bearing micritic limestones that make up the lowermost 130 m of the upper member of the Union Wash Formation and is marked by the occurrence of a 3 m-thick interval containing sphinctozoan sponges, an intervening ~1 m-thick laminated (ii = 1) micritic limestone, and an overlying, 5 m-thick bivalve-rich and bioturbated calcareous siltstone (ii = 4–5) that contains Chrondrites trace fossils. This interval is overlain by 18 m of laminated green shale (ii = 1) that signifies another incursion of anoxic waters. The third recovery interval is represented by a 30 m-thick unit that includes beds of micritic limestone, fossiliferous limestone containing transported fossil grains (including bivalves, microgastropods, sphinctozoan sponges, microgastropods, flat clams, and terebratulid brachiopods), flat pebble conglomerate and vermicular limestone. The results of this study, therefore, point to an uneven recovery within the Union Wash Formation, in which an initially robust recovery in the lower member deteriorated across the remainder of the study section. Areas exposed to persistent environmental stress, therefore, demonstrate protracted and complex recoveries. Study localities located within the habitable zone instead exhibit accelerated recoveries during the post-extinction interval and are indicative of an environmental bias in the form of persistent favorable conditions. The most accurate measures of post – extinction recovery, therefore, are those that examine biotic trends over a broad geographic area, and, as a result, incorporate the role of environmental stress in determining when and how life recovered from the Permian – Triassic mass extinction.

Introduction

The Permian-Triassic mass extinction was the most devastating biotic disturbance of the Phanerozoic (Raup 1979; Erwin 2006; Alroy et al. 2008). Recovery from the extinction has been shown to vary with regards to timing and robustness, with the amelioration of environmental conditions the main proposed facilitator of recovery (e.g., Twitchett et al. 2004; Pietsch and Bottjer 2014; Pietsch et al. 2014; Guo et al. 2015; Luo et al. 2016; Pietsch et al. 2016). Overall the absence of environmental stress appears to have accelerated biotic resurgence (e.g, Krystyn et al. 2003; Twitchett et al. 2004; Chen et al. 2007; Hofman et al. 2011; Song et al. 2011); however, incipient recoveries had the potential to be extinguished if environmental conditions worsened (e.g., Twitchett et al. 2004; Galfetti et al. 2008; Song et al. 2011). Other authors disagree about the nature of recovery, and instead argue that biotic rebound occurred independently of environmental conditions, and was instead the result of a lag between the recovery of alpha diversity (i.e., diversity of taxa) and beta diversity (i.e., differences in taxa between habitats) (Hofmann et al. 2013; Hofmann et al. 2014).
Recovering ecosystems in Early Triassic shallow shelf environments existed within a narrow “habitable zone” (Beatty et al. 2008), or “refuge zone” (Song et al. 2014) that was squeezed from below by a shallow redoxocline (Beatty et al. 2008) that capped anoxic deep waters (Isozaki 1997; Grice et al. 2005; Takahashi 2009; Song et al. 2012; Sun et al. 2015) and periodically shoaled into shallow shelf environments (e.g., Wignall and Twitchett 2002; Thomas et al. 2004; Algeo et al. 2007; Galfetti et al. 2008; Grasby and Beauchamp 2009; Bond and Wignall 2010; Liao et al. 2010). The habitable zone was also squeezed from above by hot sea surface temperatures in tropical settings that would have ranged from stressful to lethal for shallow water fauna (Romano et al. 2013; Sun et al. 2012; Song et al. 2014), as well as by wave stresses (Beatty et al. 2008). An increase in the flux of detrital material to carbonate platforms may have further stressed recovering ecosystems by increasing nutrient fluxes and the likelihood of eutrophication and by overwhelming filter-feeding organisms (Algeo and Twitchett 2010).
Western North America contains widespread Lower Triassic sedimentary rocks, and is one of the best-studied regions in terms of post-extinction recovery patterns. The earliest attempt to examine biotic recovery in western North America was by Schubert and Bottjer (1995), who document trends in macrofauna and ichnofossils from multiple localities throughout southern Nevada (Virgin Limestone), southern Utah (Virgin Limestone and Sinbad Limestone) and Idaho, Montana and northern Utah (Dinwoody Formation and Thaynes Formation). Results from this study indicate that recovery was dampened for much of the Early Triassic, and did not commence until the Anisian (early Middle Triassic) (Schubert and Bottjer 1995). The conclusions of Schubert and Bottjer (1995) have generally been supported by other studies of Lower Triassic body fossils and trace fossils from the western United States, which also find indications of a long, sluggish biotic and ecologic rebound from the mass extinction (e.g., Boyer et al. 2004; Pruss and Bottjer 2004; Nützel and Schulbert 2005; Jacobsen et al. 2011; Pietsch et al. 2014). The picture of recovery in the western United States, however, appears to be more complicated. Results from several localities demonstrate an earlier, robust recovery around the Smithian – Spathian boundary (Hofmann et al. 2013; Hofmann et al. 2014; Brayard et al. 2017), as well as earlier incipient recoveries that appear to have failed (Hofmann et al. 2014). Brayard et al. (2017) attribute this difference in recovery timing to a preservational bias or sampling errors. However, the role of deleterious environmental conditions must also be considered when documenting temporal and spatial trends in recovery (e.g., Twitchett et al. 2004; Pietsch and Bottjer 2014; Pietsch et al. 2014; Guo et al. 2015; Luo et al. 2016; Pietsch et al. 2016), since the presence or absence of environmental stress has been shown to strongly affect when recovery begins (e.g, Krystyn et al. 2003; Twitchett et al. 2004; Chen et al. 2007; Hofman et al. 2011; Song et al. 2011).

Section snippets

Geologic background

The incursion of deep, anoxic waters into continental shelf settings provides multiple, distinct sedimentologic signals (e.g., Savrda and Bottjer 1986; Wignall 1994) that can be compared to palaeontologic and ichnologic recovery trends in order to determine how biotic recovery was affected by detrimental environmental conditions. The outer shelf to slope facies of the Union Wash Formation of east-central California, U.S.A. provides a record of both biotic recovery within shallow to moderately

Methods

The Union Wash Formation was examined at the Darwin Hills, CA locality in east-central California, USA (Fig. 1a) in order to better determine the interrelationship between palaeoenvironmental conditions and recovery. A continuous section was measured across the entire 982.5 m-thick sequence. Hand samples were collected approximately every 5 m, or at greater intervals as warranted by changes in lithology; hand samples were slabbed and polished, and 125 thin sections were made from hand samples

Lower member of the Union Wash Formation

The lower member of the Union Wash Formation at the Darwin, CA locality is 152 m thick, and unconformably overlies the Lower Permian Darwin Canyon Formation (Fig. 3A). The lower member of the Union Wash Formation is characterized by 10 m-scale interbeds of brown to dark grey laminated calcisiltite and grey to brown bioturbated calcisiltite (Fig. 3B) with abundant Thalassinoides burrows that create complex and extensive networks on weathered surfaces (Figs. 3C, D). Both types of interbeds

Discussion

Three distinct recovery events are present in the Union Wash Formation at the Darwin, CA locality. Each recovery event is separated by sedimentary rocks interpreted to have been deposited under harsh environmental conditions.

Conclusions

The Lower Triassic Union Wash Formation at the Darwin, California locality contains 3 distinct spans of biotic recovery following the Permian – Triassic mass extinction that are separated by intervals of biotic stress. Recovery was strongly influenced by environmental conditions, which resulted in a recovery that was initially robust in the lower member, and deteriorated across the remainder of the unit. Overall, the results of this study indicate that areas with persistent environmental

Acknowledgments

The authors wish to thank Christopher McRoberts for help in identifying bivalves, J. single bondP. Zonneveld for help in identifying trace fossils, and Kathleen Ritterbush for useful discussions on sponges. Carlie Pietsch and one anonymous reviewer provided comments and suggestions that greatly strengthened the manuscript. Joey Hawkins provided valuable field assistance and discussions. This research was supported by funds supplied to ADW by the Department of Geological Sciences at California State

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