Shifting Mineral and Redox Controls on Carbon Cycling in Seasonally flooded Mineral Soils
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Biogeosciences, 16, 2573–2589, 2019 https://doi.org/10.5194/bg-16-2573-2019 © Author(s) 2019. This work is distributed under the Creative Commons Attribution 4.0 License. Shifting mineral and redox controls on carbon cycling in seasonally flooded mineral soils Rachelle E. LaCroix1, Malak M. Tfaily2, Menli McCreight1, Morris E. Jones1, Lesley Spokas1, and Marco Keiluweit1 1School of Earth & Sustainability and Stockbridge School of Agriculture, University of Massachusetts, Amherst, MA, USA 2Soil, Water and Environmental Science Department, University of Arizona, Tucson, AZ, USA Correspondence: Marco Keiluweit ([email protected]) and Rachelle E. LaCroix ([email protected]) Received: 4 October 2018 – Discussion started: 12 October 2018 Revised: 5 June 2019 – Accepted: 12 June 2019 – Published: 4 July 2019 Abstract. Although wetland soils represent a relatively small land soils, yet Feo had no predictive power in lowland soils. portion of the terrestrial landscape, they account for an esti- Instead, our model showed that Eh and oxalate-extractable Al mated 20 %–30 % of the global soil carbon (C) reservoir. C (Alo, a proxy of protective Al phases) became significantly stored in wetland soils that experience seasonal flooding is stronger predictors in the lowland soils. Combined, our re- likely the most vulnerable to increased severity and duration sults suggest that low redox potentials are the primary cause of droughts in response to climate change. Redox conditions, for C accumulation in seasonally flooded surface soils, likely plant root dynamics, and the abundance of protective mineral due to selective preservation of organic compounds under phases are well-established controls on soil C persistence, anaerobic conditions. In seasonally flooded subsurface soils, but their relative influence in seasonally flooded mineral soils however, C accumulation is limited due to lower C inputs is largely unknown. To address this knowledge gap, we as- through root biomass and the removal of reactive Fe phases sessed the relative importance of environmental (tempera- under reducing conditions. Our findings demonstrate that C ture, soil moisture, and redox potential) and biogeochemi- accrual in seasonally flooded mineral soil is primarily due to cal (mineral composition and root biomass) factors in con- low redox potential in the surface soil and that the lack of trolling CO2 efflux, C quantity, and organic matter compo- protective metal phases leaves these C stocks highly vulner- sition along replicated upland–lowland transitions in season- able to climate change. ally flooded mineral soils. Specifically, we contrasted min- eral soils under temperature deciduous forests in lowland po- sitions that undergo seasonal flooding with adjacent upland soils that do not, considering both surface (A) and subsurface 1 Introduction (B and C) horizons. We found the lowland soils had lower total annual CO2 efflux than the upland soils, with monthly Although wetland soils cover a relatively small portion of the CO2 efflux in lowlands most strongly correlated with redox Earth’s land surface, they store an estimated 20 %–30 % of potential (Eh). Lower CO2 efflux as compared to the up- the global soil C stocks (Mitsch et al., 2013). However, this lands corresponded to greater C content and abundance of C pool is under pressure from climate change, with increas- lignin-rich, higher-molecular-weight, chemically reduced or- ing severity and frequency of droughts having substantial, yet ganic compounds in the lowland surface soils (A horizons). largely unresolved consequences (Brooks et al., 2009; Fenner In contrast, subsurface soils in the lowland position (Cg hori- and Freeman, 2011). Increased droughts are expected to re- zons) showed lower C content than the upland positions (C lease previously stored C in wetlands back to the atmosphere horizons), coinciding with lower abundance of root biomass (Gorham et al., 1991). Prior studies focused on C cycling and oxalate-extractable Fe (Feo, a proxy for protective Fe in wetland soils have been primarily aimed at organic wet- phases). Our linear mixed-effects model showed that Feo lands, such as peats and bogs (Laine et al., 1996) or coastal served as the strongest measured predictor of C content in up- wetlands (Kirwan and Blum, 2011). Although freshwater mineral wetlands are estimated to contain 46 Pg C globally Published by Copernicus Publications on behalf of the European Geosciences Union. 2574 R. E. LaCroix et al.: Shifting mineral and redox controls (Bridgham et al., 2006), they have received comparatively anaerobic heterotrophic respiration (LaRowe and Van Cap- little attention. pellen, 2011). Anaerobic conditions limit microbes to utiliz- Previous studies on C cycling in mineral wetland soils ing substrates that are chemically more oxidized, in turn pref- have focused on permanently flooded, rather than season- erentially preserving more chemically reduced organic com- ally flooded sites (Krauss and Whitbeck, 2012). This is sur- pounds (i.e., compounds with lower oxidation states) in soils prising given that seasonally flooded soils are characterized and sediments (Boye et al., 2017; Keiluweit et al., 2017). by greater microbial activity than permanently flooded wet- While CO2 emissions are often correlated with oxygen avail- lands, resulting in significantly greater greenhouse gas emis- ability (or soil redox potential, Eh) (Koh et al., 2009), it is sions (Kifner et al., 2018). Moreover, the consequences of unclear to what extent such metabolic constraints result in climate change are expected to be most immediately evident the selective preservation of high-molecular-weight, chemi- in seasonal wetlands due to their dependence upon precipi- cally reduced OM in seasonally flooded systems where soils tation and seasonal groundwater recharge (Tiner, 2003). Sea- become aerated for prolonged periods. sonal wetlands can be considered as early warning ecosys- Water saturation also impacts soil C by controlling veg- tems (Brooks, 2005), forecasting the impacts of climate etation type and density – thus acting as an indirect con- change on permanently flooded wetlands. Thus, seasonally trol on root growth and activity belowground. Plant roots flooded wetlands represent essential end-members to study contribute to soil C stocks through rhizodeposition (exu- the effects of climate change on permanently flooded wet- dates, secretions, dead border cells, and mucilage), dead root land soils (Brooks, 2005). residues (Jones et al., 2009), and root-associated microbes Seasonal wetlands are geomorphic depressions in the (Bradford et al., 2013). Roots are the main contributors to landscape that have distinct hydrologic phases of flooding C stocks in upland soils (Rasse et al., 2005), but the im- and draining (Brooks, 2005). These ephemeral wetlands are pacts of roots on soil C stocks in wetlands are less clear. small (< 1 ha) but ubiquitous – comprising nearly 70 % of Water saturation directly inhibits root growth due to the asso- all temperate forest wetlands in the US (Tiner, 2003). Sea- ciated low dissolved oxygen concentrations (Day and Mego- sonal flooding and drainage not only create biogeochemi- nigal, 1993; Tokarz and Urban, 2015). Indirectly, water satu- cal “hotspots” for soil C and nutrient cycling along upland– ration in soil selects for plant species that can tolerate water lowland transitions but also “hot moments” as these tran- stress – typically species that have developed advantageous sition zones move seasonally (Cohen et al., 2016). These traits to survive flooded conditions, such as shallow root- transition zones are also relatively large, as the generally ing systems (Tokarz and Urban, 2015). However, seasonally small size of seasonal wetlands results in a disproportion- flooded soils select for an even smaller niche of plants, as ally large and dynamic terrestrial–aquatic interface relative they must be tolerant of both upland and lowland conditions to total wetland area (Cohen et al., 2016). Determining the (Brooks, 2005). How root inputs from facultative upland– controls on C cycling within seasonally flooded mineral soils lowland plant species contribute to soil C content and chem- thus requires specific consideration of the fluxes and dynam- istry in seasonally flooded soils is still not clear. ics across these terrestrial–aquatic transitions. In addition to restricting microbial metabolism and root Temperature and soil moisture are principal controls on C growth, water saturation influences the concentration and cycling in soils (Lloyd and Taylor, 1994; Wang et al., 2014). distribution of high surface area minerals that are potent sor- Temperature regulates biological and chemical reaction rates bents for C in soils (Chen et al., 2017; Torn et al., 1997; and thus regulates the rate at which decomposition of soil or- Wagai and Mayer, 2007). In upland soils, iron (Fe) or alu- ganic matter can occur (Davidson and Janssens, 2006). How- minum (Al) (hydr)oxides protect OM from microbial decom- ever, water saturation is a critical driver of organic matter position, thereby contributing to C storage for centuries to (OM) decomposition processes in seasonally flooded sys- millennia (Torn et al., 1997; Wagai and Mayer, 2007). In tems (Neckles and Niell, 1994). Water saturation governs flooded soils, however, the rapid depletion of oxygen upon oxygen availability in soil pore spaces, as oxygen diffusion flooding can result in the reductive dissolution of Fe(III) ox- in water is 10 000 times slower than in air (Letey and Stolzy, ides (Chen