Revisiting carbon cycling in the Laurentian Great Lakes following dreissenid mussel invasion

doi:10.21203/rs.3.rs-4436844/v1

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Revisiting carbon cycling in the Laurentian Great Lakes following dreissenid mussel invasion

https://doi.org/10.21203/rs.3.rs-4436844/v1

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Since the active role of inland waters in cycling carbon (C) has been revealed, there has been a renewed interest in calculating C budgets for inland waters to understand their role with respect to global climate change. There is a lack of knowledge regarding C cycling in the Laurentian Great Lakes, the worlds largest freshwater reservoir, with current estimates neglecting the role of invasive species. For one of the most pervasive invaders, dreissenid (zebra and quagga) mussels, research has focused on filter feeding impacts on phosphorus dynamics, but there is a lack of knowledge regarding their role in C cycling, specifically, the impact of the C stored in their slowly degrading shells. As such, we set out to estimate the mass of empty shells and C stored in those shells. We calculated an estimated 1.19 E10 tonnes of empty shell mass currently sitting at the bottom of these lakes, which store approximately 1.43 E9 tonnes of C. This scale of inorganic C storage is comparable to rates of organic C storage in nature-based climate solutions. This work demonstrates the importance of a previously unexplored pathway that dreissenid mussels are altering C cycling in the Laurentian Great Lakes and the thousands of other invaded lakes and rivers.

Earth and environmental sciences/Environmental sciences

Earth and environmental sciences/Limnology

Earth and environmental sciences/Climate sciences/Biogeochemistry

carbon cycling

dreissenid mussels

Laurentian Great Lakes

inland carbon budgets

mussel shells

inorganic carbon deposition

The calculation of up-to-date, accurate carbon (C) budgets is essential for climate change mitigation and adaptation plans ^1,2. There has been a large amount of work developing C budgets, especially in terrestrial and oceanic environments, however, more work is required to understand the C budgets of inland waters ^1,3,4. Inland waters have previously been considered as passive ‘pipes’ transporting terrestrial C to the oceans, however, recently there has been work showing the active role of inland waters in global C cycling ^3,5–7. Globally, the majority of C in inland waters (streams, rivers, lakes, reservoirs, and ponds) is from terrestrial sources (5.8 Pg C/yr; inorganic and organic C) with a small amount contributed from aquatic primary productivity (0.3 Pg C/yr; organic C), of this C supply, 4.4 Pg C/yr is emitted to the atmosphere, 1.1 Pg C/yr is exported to the oceans, while just 0.6 Pg C/yr is buried in aquatic sediments ⁷.

Although the annual rate of carbon burial in lakes is a relatively small, sedimentation in inland waters presents a long-term carbon storage pathway ^3,6,8. Lake size and primary productivity have been identified as key factors determining sedimentation rates, in addition, cultural eutrophication has been shown to increase C sedimentation rates by increasing primary productivity ^8–11. ^{[NO_PRINTED_FORM] 11}, estimated that European lakes had an average pre-industrial organic C burial rate of 5–10 g C/m²/yr, which increased to an average of 60 g C/m²/yr post-1950. In North America, ^{[NO_PRINTED_FORM] 10} calculated an average organic C burial rate of 88 g C/m²/yr across eight lakes in the Midwestern region of the United States. In the Great Lakes, Lake Erie and Ontario have higher primary productivity compared to the upper lakes (Huron, Superior, and Michigan), and organic C burial rates mirror this trend (19.07–30 vs. 0.97–4.84 g C/m²/yr), despite the smaller size of the downstream lakes ¹². Additionally, in the Great Lakes, dreissenid mussel filtration has been explored as a factor influencing C sedimentation as it consumes large amounts of algae and takes up calcium for shell growth, reducing its concentrations for precipitation ^13–16.

While organic C burial is increasingly well resolved in freshwater ecosystems, the inorganic C cycle of inland waters has not been as extensively explored, despite the important role that inorganic C plays in inland water C cycling ^17–20. In marine systems, a study on high Arctic fjords found inorganic C burial rates ranged between 10.7–45.7 g C/m²/yr, with inorganic C burial dominating organic C at one of the two fjords sampled ²¹. ^{[NO_PRINTED_FORM] 21}, attributed the dominance of inorganic C burial to a greater activity of calciferous organisms and spread of carbonates in the area. The role of inorganic C burial has also been investigated for coastal blue carbon (CO₂ stored in coastal saltwater environments) and global inorganic C burial rates were estimated at 0.8 Tg C/yr for mangrove ecosystems and 15–62 Tg C/yr in seagrass ecosystems ²². However, when determining the impact of inorganic C in bottom sediments on C budgets, it is essential to consider the C source. Allochthonous inorganic C originates from terrestrial respiration and is transported to aquatic ecosystems (via runoff and groundwater), where it may be taken up for primary production or degassed to the atmosphere ^3,22. Counterintuitively, the burial of inorganic C via mineralization increases CO₂ concentrations as CO₂ is a byproduct of CaCO₃ mineralization (Eq. 1). Additionally, the reverse reaction, dissolution of CaCO₃, results in an uptake in CO₂. Consequently, the net inorganic C burial from mineralization offsets the CO₂ sink from organic C burial, in the case of the blue carbon ecosystems described above, a ~ 30% offset was calculated ²². Thus, it is evident that omitting inorganic C burial in inland water C budgets can result in inaccurate carbon pool evaluations, especially in systems associated with high mineralization from calciferous organisms ^3,23. Although the above examples provide insights on inorganic C burial from calciferous organisms on marine C budgets, there is a gap in their consideration for freshwater environments ^7,12,22.

Although the indirect impacts of dreissenid filter feeding have been briefly explored in relation to organic C sedimentation ¹³, the impact of shell calcification and burial has not previously been considered for inland C cycling ^14,15.

Equation 1. ${2HCO}_{3}^{-}+{Ca}^{2+}{\leftrightarrow CaCO}_{3}+{CO}_{2}+{H}_{2}O$

In the above stated studies inorganic C from calcification was considered a source of CO₂ to the environment, however this may not be entirely accurate as it depends on the C source for calcification ^24–26. Previously, dissolved inorganic carbon (DIC) was considered the sole source of mussel shell C and thus that C isotopes in shell were thought to directly reflect their environmental conditions, just as stable isotope analysis of oxygen in shells has been used to reflect ambient temperature throughout a mussel’s lifetime ²⁷. More recently, it has become clear that metabolic C can make up a notable proportion of mussel shell C (%C_M), ~ 12% in marine mussels but up to 25–40% in freshwater mussels ^25,26,28,29. Specifically, a study of the North American freshwater mussel, Elliptio complanata, found an average %C_M of 20 ± 10%, the large uncertainty represents the variation due to factors including ontogenic shifts, metabolic rates, and food source and selectivity ²⁸. Determining the relative proportion of C sources is not only important for paleo work, but also to understand the impact of modern mineralization on C cycling. While shell mineralization from DIC results in the release of a CO₂ molecule, when organic C is the original source a molecule of CO₂ from respiration is diverted from the release to the environment. Thus, the relative proportion of C sources for shell growth in dreissenid mussels need to be determined to understand their impact on inland C cycling (Fig. 1).

The Laurentian Great Lakes (hereafter Great Lakes) contain ~ 18% of the world’s liquid, surficial freshwater, with a watershed of approximately 1.0 million km^{2 30}. Due to cultural eutrophication, most of the budget calculations in the Great Lakes focused on nutrients (nitrogen and phosphorus) rather than on carbon. Preliminary calculation indicates that the carbon budget is dominated by autochthonous production, followed by inorganic and organic C inputs from stream and groundwater and surface runoff ¹². Previous work has also identified the Great Lakes as a net source of CO₂ to the atmosphere, although CO₂ saturation decreases from the upstream to downstream lakes, and varies seasonally ^13,31–33. Additionally, ^{[NO_PRINTED_FORM] 13}, found an increase in Great Lakes pCO₂ saturation following the establishment of large quagga mussel (Dreissena bugensis) populations and hypothesized that this was due to the alteration of lake water biogeochemistry via decreased primary productivity, increased water clarity, and respiration from the large mussel populations. This finding draws from a large body of work examining the physicochemical impacts of invasive dreissenid mussels ^34,35, especially in the Great Lakes ^36–40. Although the impacts of living dreissenid mussels are well understood in the Great Lakes and other invaded territories, there has been very little work examining the impacts of the accumulating mass of mussel shells on the carbon budget in these lakes ^38,41,42.

There has been some work exploring the scale and role of dreissenid mussel shells in inland lakes, however, as of yet, the Great Lakes, the largest freshwater ecosystem on Earth, have not been considered in this field. In Europe, a study investigated the impact of dreissenid mussel shell accumulation on a shallow lakes’ life span, and found that the invasion of D. bugensis could shorten the life span of the lake by one- to two-thirds by filling it with spent shells ⁴³. Additionally, ^{[NO_PRINTED_FORM] 44}, investigated shell decay rates for native and invasive bivalves in North American waters and found that Dreissena spp. can produce standing stocks of spent shells on the scale of > 10 kg dry mass/m² under appropriate conditions (low flow, high calcium) ⁴⁴. ^{[NO_PRINTED_FORM] 45}, collected benthic samples from ten Minnesota lakes and although they found high variability in empty shell mass across lakes, they determined that dead shell material accounted for an average of 322% of lake particulate organic carbon (POC). Finally, the most comprehensive study of dreissenid shell accumulation in a large lake, was conducted in Lake Simcoe, Ontario, Canada (SA = 722 km²) ⁴⁶. Based on samples collected from 0–32 m depths, a total dead shell mass of 1 141 000 tonnes was estimated ⁴⁶. Additionally, the authors found that approximately 1 000 000 tonnes of shells are produced annually in Lake Simcoe, containing ~ 115 600 tonnes of C.

Since their invasion of the Great Lakes in the late 1980s, dreissenid mussel populations have had consistent growth, with an initial growth of the zebra mussel (D. polymorpha) population followed by the delayed establishment of quagga mussels ^47–49. The establishment of quagga mussels not only increased dreissenid density in previously established zebra mussel habitat, but also expanded to much greater depths (> 90 m) and softer substrates ^47,50,51. The growing dreissenid mussel population was also able to withstand predation pressure from waterfowl, lake whitefish, and their native predator, the round goby (Neogobius melanostomus), which established Great Lake populations in the early 2000s ^40,50,52. Predation rates vary by habitat, but previous work has estimated round goby predation of dreissenid mussels in Lake Erie to be six kT/yr (270 T C_shell/yr), and lake whitefish consumption of 109 and 820 kT/yr (500 and 3800 T C_shell/yr) in Lakes Michigan and Huron, respectively ^40,53.

While dreissenid predation has increased with long-term establishment, there is minimal evidence that these predators are reducing dreissenid abundance, although a shift in size structure has been observed ^52,54,55. Predation rate and the dreissenid population impact the turnover rate of dreissenid mussels (and production of empty mussel shells), through their impact on the production: biomass (P:B) ratio. The stable dreissenid population maintains biomass (B), while the increased predation rate results in increased dreissenid production (P) (Astorg et al., 2022; Kipp & Ricciardi, 2012; Naddafi & Rudstam, 2014). Therefore, the combination of stable dreissenid populations with increased predation suggests that dreissenid mussel turnover has increased, thus increasing the rate of dreissenid shell mineralization in the Great Lakes (Fig. 2).

The evidence of large-scale inorganic C sequestration in much smaller North American lakes and a large stable dreissenid population in the Great Lakes makes a clear case for calculating C sequestered in empty mussel shells in the Great Lakes. Altogether it is evident that dreissenids are persistent ecosystem engineers in the Great Lakes ^34,36,50, however, the impact of nutrient sequestration in their shells that are continuously accumulating at the bottom of these lakes has not previously been considered as a component of their engineering role. Here we provide an estimation of the C mass in empty dreissenid shells in the four dreissenid-invaded Great Lakes (Michigan, Huron, Erie, and Ontario). This quantification provides a starting point for considering another pathway that dreissenid mussels have altered C dynamics in the Great Lakes. Finally, we suggest future work that will better resolve the impacts of this pathway on C budgets.

To estimate the scale of mineralized C in empty dreissenid mussels in the Great Lakes we calculated the empty shell standing stock based on average pre- (1990–2005) and post (2005–2020)-goby invasion mussel densities from 1990–2020 (Table 2). Estimates were split by pre- and post-goby periods to account for the increased dreissenid mussel mortality (Z) following the establishment of stable round goby populations ^52–54. Round goby invasion also coincided with quagga mussel dominance which occurred in most lakes by the year 2000 and expanded the mussel population to deeper depths ^51,56. This combination of increased mussel abundance (biomass) and predation pressure (mortality rate) resulted in much higher shell turnover and thus higher mineralized C in empty shells (2.40 X 10⁸ vs 1.19 X 10⁹ T C). This shift was most prominent in Lakes Ontario and Michigan, where increased mussel densities in the deeper waters and increased predation resulted in much higher C mineralization in the post-round goby period ^42,47,57. An estimated total of 1.19 E10 tonnes of empty shell mass was calculated across the four lakes, containing 1.43 E9 tonnes of C.

Table 2

Estimated carbon stored in empty dreissenid shells (in tonnes of carbon) in the Great Lakes before and after the invasion of its native predator, the round goby. Based on average 15 years pre-goby invasion (1990–2005) and 15 years post-goby invasion (2005–2020) mussel densities.
	Lake Ontario	Lake Erie	Lake Huron	Lake Michigan	Total
Pre-round goby C storage (T C)	1.15 X 10⁷	1.74 X 10⁸	1.64 X 10⁷	3.80 X 10⁷	2.40 X 10⁸
Post-round goby C storage (T C)	1.11 X 10⁸	9.02 X 10⁷	9.83 X 10⁷	8.92 X 10⁸	1.19 X 10⁹
Total C storage (T C)	1.23 X 10⁸	2.64 X 10⁸	1.15 X 10⁸	9.30 X 10⁹	1.43 X 10⁹

Dreissenid mussel shells clearly represent a large deviation in carbon flow within the lakes and need to be considered in future carbon budget calculations ¹². This work is a key step in filling the knowledge gap regarding the role of aquatic ecosystems as contributors or sinks of carbon to the atmosphere. ^3,12,58. Dreissenid mussels have existed in the Great Lakes for just over 30 years and our calculations indicate that their accumulated spent shells account for 1.43 Gt of inorganic C. Throughout time, Canadian lakes are estimate to have stored a total of 20 Gt of C, our calculations indicate that inorganic C from dreissenid shells would make up ~ 7% of this stored C ^58,59. On an areal basis, there was a large range of inorganic C deposition rates across the lakes (0.08–37.5 T C/ha/yr), which were primarily dependent on mussel density. This range of C storage is on the scale of estimates of organic C storage from nature-based climate solutions (NBCSs) and depending on the C source (metabolic C or DIC), may supplement or offset these NBCS’s C storage (Fig. 3). Based on the current literature, it is likely that DIC is the primary source, indicating that dreissenid shell mineralization will release CO₂ to the environment, offsetting the CO₂ sink from NBCS’s ^26,28. Dreissenid mussel inorganic C deposition is also on a similar scale to organic C sequestration rates in established coastal blue ecosystems (seagrass, mangroves, and salt marshes) (Fig. 3; Alongi, 2023; Claes et al., 2022). When compared to Canadian freshwater NBCSs, inorganic C is deposited from dreissenid mussel shells in the Great Lakes at a much greater rate than organic C in wetlands and peatlands. Specifically, the inorganic C from shells is deposited at a rate greater than organic C stored in peatlands (23 g/m2/yr), freshwater marshes in the Lake Simcoe watershed (average 112 g C/m²/yr), and other intact (62 g C/m²/yr) and restored (51–89 g C/m²/yr) wetlands in southern Ontario ^62,63.

The inorganic C deposition of dreissenid shells is also on the scale of inorganic carbon storage calculated in other systems. ^{[NO_PRINTED_FORM] 21}, found inorganic C burial in Arctic fjords at a rate of 10.7–45.7 g C/m²/yr, which is within the range of our rate of ~ 0.2–3750 g C/m²/yr, with a median of 201 g C/m²/yr. On a global scale, seagrass ecosystems have been found to accumulate inorganic C at rates from 149–756 g C/m²/yr ^22,64–66. However, it is important to note that these accumulation rates are based on all inorganic C deposited in bottom sediments (allochthonous and autochthonous) and not specific to calcifying organisms ²². When looking specifically at inorganic C accumulation from local calcification in seagrass, the range is much lower, 2.2–36 g C/m²/yr ^64–67. Thus, the inorganic C accumulation from dreissenid mussels in the Great Lakes is at a scale that should be considered in future C sequestration rate and C budget calculations.

The estimates provided here are based on historical turnover and shell degradation rates, however, it is essential to consider the impact of predicted future changes to the Great Lakes on dreissenid shell C burial. An updated investigation into shell decay rates in the Great Lakes is recommended to provide a more accurate value for future calculations. ⁴⁴ found that shell decay rates were correlated with pH and flow, however D. bugensis occupy much deeper habitats than D. polymorpha and other depth related variables (DO, light, nutrients etc.) should be considered for future calculations, especially as climate change is expected to change these conditions ⁶⁸. Additionally, winter ice scour can result in high over-winter mussel mortality at shallow depths ⁶⁹. Climate change is predicted to alter the ice-cover period in the Great Lakes ^70,71, which could reduce the occurrence of ice scour mortality. Future changes to other factors that affect mussel population dynamics (nutrient levels, substrate) should also be considered since they will influence the rate of shell deposition as well. Following their initial establishment, dreissenid populations were relatively consistent in their invaded territories, even following the establishment of their native predator, the round goby ^42,72. However, it is possible that new predators may be introduced, or food source availability may change, altering mussel mortality rate ^70,71

Dreissenid mussels are considered one of the most impactful invasive species in North American waters. Serving as ecosystem engineers, they modify benthic habitats and influence phosphorus cycling and nuisance algae growth ^34,35,70. We present here another ecosystem-scale impact that expands their role in nutrient cycling to include inorganic C sequestration. On the scale of billions of tonnes of inorganic carbon deposited in slowly degrading shells at the bottom of these lakes. Although our calculations provide an estimate of the scale of their C deposition capacity, physically measuring the mass of shells in the Great Lakes and connecting channels, as ^{[NO_PRINTED_FORM] 46} did in Lake Simcoe, will provide the best estimation C stored in empty dreissenid shells. An essential part of this work will be determining the relative proportions of each shell C source (metabolic and DIC), this will provide the insight needed to determine the role of dreissenids on C cycling and CO₂ flux in the Great Lakes and other invaded territories.

The storage of other elements, such as nitrogen and phosphorus, in dreissenid mussel shells should also be calculated. Although the proportions of shell nitrogen (~ 0.25%) and phosphorus (~ 0.007%) are much smaller than carbon (~ 12%) ^37,45,46, based on the shear scale of deposited shells they may be an important sink for these nutrients as well, especially when considering that phosphorus concentrations have declined in most of these lakes ^70,73. Finally, the creation of shell fragments from predation needs to be considered. Previous calculations of shell decay rates are based on whole shells suspended in the water column, however, most dreissenid mussel predators break the shells into fragments to consume the mussel tissue and discard shell fragments that settle into the sediments ⁵⁴. As described in the Methods, it is not clear how these small shell fragments will impact dissolution rates, they could reduce dissolution by increasing shell compaction (reducing exposed surface area) or as smaller particles they could dissolve more quickly ⁴⁴. If dissolution does increase it would result in CO₂ uptake from the water column through the reverse of Eq. 1. Evidently, determining the rate of shell dissolution with shell fragments will provide a better understanding of dreissenid mussel impacts on C cycling in the Great Lakes.

This work adds to the previously explored roles of dreissenid mussels as ecosystem engineers in the Great Lakes and other invaded waterways. In addition to their phosphorus cycling role and increased respiration rates, dreissenid mussels are altering the flow of carbon through mineralization on a massive scale ^13,35,36. As mentioned above, determining the relative role of metabolic C vs. DIC in shell mineralization is key to deciphering their exact impact on C cycling and CO₂ flux in invaded waters. Previous work indicates that ambient bicarbonate is likely the primary source, and thus until the relative proportions are determined we should consider dreissenid shell mineralization as a likely source of CO₂ to invaded waters ²⁸. While dreissenid mussels have been found to alter CO₂ concentrations through respiration and impacts on primary productivity, their shell mineralization has not been considered as a component of this impact ^13,74. Regardless of the direction (sink or source) it is evident that shell mineralization has major impacts on C cycling in the Great Lakes.

On a larger scale these findings have implications for calculation of C budgets for the Great Lakes and implications for the global inland waters C balance. A preliminary C budget for the Great Lakes indicates that net primary production (NPP) and respiration are the greatest C flows for all lakes except Ontario, which receives more C from upstream inputs ¹². Our calculations indicate that dreissenid mineralization alters C flow on the same scale as the processes considered in ¹² calculations and should be included in future C budgets for the invaded lakes (Michigan, Huron, Erie, and Ontario). Additionally, these C fluxes should be included in global inland water C budgets, although we focused on the Great Lakes, there are thousands of invaded lakes and rivers across North America and Europe that will have altered C flows due to dreissenid mussel mineralization ^34,35. Finally, inclusion in these C budgets will have policy implications in the corresponding jurisdictions. Many locations have drafted adaptive management plans to minimize greenhouse gas emissions and an improved understanding of local C dynamics will make these plans more robust as dreissenid mussel populations continue to grow.

Calculation of dreissenid empty shell standing stock was based on ⁴⁴, where at steady state the standing stock of spent shells (SS) is equal to the production of spent shells (P) divided by the instantaneous decay rate (I). Production of spent shells was determined from living shell mass (B) and instantaneous mortality rate (Z, or P:B depending on availability), with Eq. 1.

Equation 1.

$$SS=\frac{B*Z}{I}$$

The instantaneous shell decay rate of 0.058 calculated by ⁴⁶ was used as it is comparable to other estimates ⁴⁴, and was calculated in Lake Simcoe, a lake in the same region as the Laurentian Great Lakes, with similar geochemical characteristics. This decay rate is a very conservative estimate as it is based on shells suspended in the water column, where turbulence is greater than on the lake bottom, which would result in an overestimation of the dissolution rate. Additionally, the settlement of shells in layers on the lake bottom would reduce exposed surface area and likely result in slower decay compared to shells in the water column. Finally, the increased shell fragmentation from round goby predation could impact shell decay rate, however without published findings on this impact it is not possible to adjust decay rate for this factor. Shell fragmentation would alter the matrix of deposited shells potentially reducing decay rate by reducing exposed surface area, alternatively ⁴⁴, showed that smaller shells result in higher decay rates. Considering this contradiction, the published value was used to avoid making an incorrect assumption due to the lack of literature on this factor.

Two mortality rates were used, one for the pre-round goby period (~ 1990–2005) and one for the post-round goby period (~ 2005–2020). The pre-round goby, non-predatory mortality rate of 1.26/year from ⁷⁵. The post-round goby production: biomass ratio (P: B)/mortality rate of 2.1 from ⁷⁶ was applied for the post-round goby invasion period ~ 2005–2020.

An average living shell biomass was determined for each period based on published density values for each depth zone (< 30 m, 31–50 m, 51–90 m) for Lake Ontario, Lake Huron, and Lake Michigan. As Lake Erie is very shallow and has more variation across its basins (East, Central and West) than depth, shell biomass was determined from mean basin-wide biomass. Lake Superior was not included in calculations as it’s dreissenid populations are limited to select areas of the lake, and extrapolating to a lake-wide estimation based on depth zones would not be appropriate ^50,77. Living dreissenid density data was extracted from lake-wide surveys with at least 30 sites, across a range of bottom substrates to represent the range of conditions in each lake (Table 1). Additionally, where a series of similar surveys were conducted the most recent published data was extracted. All surveys collected between 2–6 replicates for each sample site.

Table 1

Summary of dreissenid density data sources for each lake and period (pre- and post-round goby invasion).
Lake	Period	Source	Year(s) sampled	Sample sites (#)	Depth range (m)
Lake Ontario	Pre-goby	⁸³	1997	75	14–209
Lake Ontario	Post-goby	⁴²	2008, 2013, 2018	55	11–209
Lake Erie	Pre-goby	⁷⁸	1998	30	2.0–58.5
Lake Erie	Post-goby	⁸⁴	2009, 2012	135	2–66
Lake Huron	Pre-goby	⁴⁸	2000, 2003	65–85	19–173
Lake Huron	Post-goby	⁴⁸	2007, 2012	80–83	12–171
Lake Michigan	Pre-goby	⁴⁹	1994,1995,2000	90–157	6–208
Lake Michigan	Post-goby	⁴⁹	2005, 2010	140–160	13–207

Living mussel shell density was calculated from the wet mass density (g/m²) assuming shells are approximately 38.13% of whole mussel wet weight (Báldi et al., 2019; Table S4), if whole mussel wet weight was not available it was calculated based on density of individuals (ind./m²) using an average wet weight of 0.156 g/ind. ^16,78. Empty shell standing stock was multiplied by 0.12, the approximate proportion of carbon in the shell ^37,45, to get the areal mass of carbon in empty shells deposited each year. Lake surface area for each depth zone described above was calculated from NOAA bathymetry maps ^79–82. Standing stock of empty dreissenid shell C deposited per year was calculated by combining areal production with each zone’s surface area. Finally, the total mass of C stored in empty dreissenid shells was calculated by multiplying the pre- and post-round goby estimates by 15 years, the approximate time for each period.

Competing Interests

The authors declare no competing interests.

Author Contribution

EDS: Data curation, formal analysis, writing – original draft preparation. LJM: Conceptualization, supervision, writing – review & editing. JM: Writing – review & editing. AEK: Writing – review & editing. JR: Supervision, Writing – review & editing.

Acknowledgement

We would like to thank all members of our team for their efforts. The conceptual models were created by Lexy Harquail. We would also like to thank our funding agencies: Mitacs and RBC Foundation.

Data Availability

All data generated or analysed during this study are included in this published article [and its supplementary information files].

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Revisiting carbon cycling in the Laurentian Great Lakes following dreissenid mussel invasion

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