the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Reviews and syntheses: Bioturbation impacts on sediment accretion and erosion in tidal marshes, with implications for carbon burial and sequestration
Abstract. Tidal marshes offer multiple ecosystem services, but are some of the most threatened coastal ecosystems worldwide. One of these valued services is their ability to sequester and store large amounts of carbon. Bioturbating macrofauna are ecosystem engineers that can influence the geomorphology and biogeochemistry of tidal marshes. Bioturbators can influence accretion and erosion processes in tidal marshes by either stabilizing or destabilizing sediment. Through this reworking of sediment, they can also influence the amount of carbon that can be stored. The impact of bioturbation on tidal marshes depend on a number of factors, such as, species composition, burrow morphology, diet, behaviour and habitat type. This review assesses the current knowledge on the role benthic bioturbators play in shaping sediment processes in tidal marshes and identifies key knowledge gaps for future research. For example, the impact of individual benthic species on sediment dynamics is mostly unknown. Bioturbation effects cannot be generalised and predicting when and where these effects will be most prominent is challenging. Future studies should investigate family and species specific effects on sediment properties, such as erodibility or texture, under controlled laboratory conditions and in the field. This should be compared across different habitat types such as ecotones, mudflats, salt marshes and mangroves. Furthermore, the role of consumers, as bioturbators, remains an understudied driver of the carbon cycle because it is complex. In order to better predict how tidal marshes may persist in the face of future climate change, such as sea level rise, it is important to understand the role of bioturbators on sediment and carbon dynamics to enable better mitigation of global change effects through conservation and restoration of tidal habitats.
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RC1: 'Comment on egusphere-2025-4749', Anonymous Referee #1, 12 Nov 2025
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AC1: 'Reply on Reviewer 1's general comment', Leigh-Ann Smit, 02 Feb 2026
This manuscript presents an overview of the effect of bioturbation on tidal wetlands. While the topic is of great interest, the material is poorly organized, making it difficult for readers to develop a general understanding of the subject. As it stands, the manuscript reads more like a collection of disparate results from the literature rather than a cohesive synthesis. Below, I outline major points for improvement, along with specific suggested edits.
Thank you for this feedback and for the useful comments provided, which we have addressed below. We will especially focus on restructuring the text to ensure flow and coherency.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC1 -
AC2: 'Reply on RC1', Leigh-Ann Smit, 02 Feb 2026
An overview of different bioturbators across continents and regions should be provided early in the paper.
We agree that adding this information early on in the paper would improve the contextual framing of the review. We will add this to section 1.3 (Bioturbation in coastal wetlands). This section introduces the organisms associated with bioturbation, so we will incorporate the types of bioturbators and where they are found. This will highlight the typical organisms (down to species-level where information is available) found across the tidal wetlands of global bioregions. A new table will be included to summarise the information.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC2 -
AC3: 'Reply on RC2', Leigh-Ann Smit, 02 Feb 2026
A detailed description of the physics of burrowing, and how it varies depending on species, substrate, and region, is currently missing and should be included.
We will expand the manuscript to provide this information as suggested. We will include physical mechanisms of burrow construction and maintenance, looking at compaction, bioirrigation and sediment displacement. This will also include how these processes differ across different taxa and substrates. This will be added to section 1.3 (Bioturbation in coastal tidal wetlands). A new table will be included to summarise this information.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC3 -
AC4: 'Reply on RC3', Leigh-Ann Smit, 02 Feb 2026
The introduction is fragmented and contains much extraneous information; it needs to be more focused on the central topic.
We will reorganize and streamline the introduction by adding subheadings and removing unnecessary information. We will include headings such as “An overview of tidal wetlands”, “Key sediment processes” and “Key carbon processes”, as suggested by reviewer 2 (comment 2.1).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC4 -
AC5: 'Reply on RC4', Leigh-Ann Smit, 02 Feb 2026
Only two figures are presented, which is insufficient. The manuscript would benefit from additional conceptual diagrams illustrating main processes and feedbacks. Figures that summarize the text would be particularly helpful.
Although Figure 3 summarises many of the processes influenced by bioturbators, we will add additional illustrations to improve clarity and highlight key processes, especially in line with comment 1.2 above to summarise these. We will add the dominant regional bioturbators to Figure 2 and create an additional figure on the burrowing modes and effects on sediment of different bioturbators (e.g. crab burrows, worm burrows, shrimp burrows). We will also include a conceptual summary figure at the end of the manuscript with the major feedbacks, both positive and negative and link it to sediment dynamics and carbon processes.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC5 -
AC6: 'Reply on RC5', Leigh-Ann Smit, 02 Feb 2026
Including tables that compile data from multiple studies to generalize results would greatly strengthen the manuscript.
Table S1 provides a synthesis of the literature and studies provided in the text. We will draft two summary data tables in response to comments 1.1 and 1.2 above, which will also thereby summarise the information presented in Table S1.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC6 -
AC7: 'Reply on RC6', Leigh-Ann Smit, 02 Feb 2026
Including tables that compile data from multiple studies to generalize results would greatly strengthen the manuscript.
Table S1 provides a synthesis of the literature and studies provided in the text. We will draft two summary data tables in response to comments 1.1 and 1.2 above, which will also thereby summarise the information presented in Table S1.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC7 -
AC8: 'Reply on RC7', Leigh-Ann Smit, 02 Feb 2026
A dedicated section or paragraph on species migration driven by global warming is missing and should be incorporated.
We will add an additional paragraph to section 1.6 (Global change impacts on tidal marsh bioturbation), to include this information. We will focus on projected shifts in the distribution of species and how this might influence bioturbation patterns, with implications for sediment and carbon processes under these scenarios.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC8 -
AC9: 'Reply on RC7', Leigh-Ann Smit, 02 Feb 2026
Some additional minor edits are noted in the text.
We note the very useful 54 edits and comments made by this reviewer throughout the PDF manuscript that assist us in addressing the seven broad comments above, and have prepared the following responses to these:
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC9 -
AC10: 'Reply on RC8.1', Leigh-Ann Smit, 02 Feb 2026
Line 2-Replace “marshes” with “wetlands”.
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC10 -
AC11: 'Reply on RC8.2', Leigh-Ann Smit, 02 Feb 2026
Line 17- Replace “influence” with “affect”.
This will be changed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC11 -
AC12: 'Reply on RC8.3', Leigh-Ann Smit, 02 Feb 2026
Line 37- The introduction is too fragmented with too much information, you might want to reduce it and make it more focused
We will reorganise the text and include headings such as “An overview of tidal wetlands”, “Key sediment processes” and “Key carbon processes”. We will also remove any unnecessary information.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC12 -
AC13: 'Reply on RC8.4', Leigh-Ann Smit, 02 Feb 2026
Line 39- Replace “marshes” with “wetlands”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC13 -
AC14: 'Reply on RC8.5', Leigh-Ann Smit, 02 Feb 2026
Line 40- Change “exist between” to “connect”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC14 -
AC15: 'Reply on RC8.6', Leigh-Ann Smit, 02 Feb 2026
Line 43- Replace “they” with “Tidal wetlands”
This will be changed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC15 -
AC16: 'Reply on RC8.7', Leigh-Ann Smit, 02 Feb 2026
Line 50- Replace “with” with “and”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC16 -
AC17: 'Reply on RC8.8', Leigh-Ann Smit, 02 Feb 2026
Line 55- Replace “their” with “its”
This will be replaced.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC17 -
AC18: 'Reply on RC8.9', Leigh-Ann Smit, 02 Feb 2026
Line 57- Include a space between the words “abiotic” and “and”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC18 -
AC19: 'Reply on RC8.10', Leigh-Ann Smit, 02 Feb 2026
Line 57- Remove “s” at the end of includes
We will change it to “include”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC19 -
AC20: 'Reply on RC8.11', Leigh-Ann Smit, 02 Feb 2026
Line 62- Replace “loses” with “losses”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC20 -
AC21: 'Reply on RC8.12', Leigh-Ann Smit, 02 Feb 2026
Line 62- Change “estimated” to “assessed”
This will be changed
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC21 -
AC22: 'Reply on RC8.13', Leigh-Ann Smit, 02 Feb 2026
Line 62- Reference needed
The reference “Mentaschi et al. 2018” will be added.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC22 -
AC23: 'Reply on RC8.14', Leigh-Ann Smit, 02 Feb 2026
Line 63- This paragraph “Some studies have revealed that accretion rates are insufficient for tidal marshes to keep pace with sea level rise (e.g. Van Wijnen and Bakker, 2001), while others have found that accretion rates are high enough to keep pace with moderate rises in sea level (e.g. Morris et al., 2002)” is out of context
This section will be moved to section 1.6 (Global change impacts on tidal marsh bioturbation).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC23 -
AC24: 'Reply on RC8.15', Leigh-Ann Smit, 02 Feb 2026
Line 82- Provide clarity on what is meant by relative to the ocean carbon cycle
This will be made clearer in the text as follows “These blue carbon habitats store up to 70% of carbon, relative to the ocean carbon cycle, despite covering less than 2% of the area of the global ocean”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC24 -
AC25: 'Reply on RC8.16', Leigh-Ann Smit, 02 Feb 2026
Line 96- What is the difference between high productivity and conversion of carbon dioxide into plant biomass
High productivity is referring to how much biomass accumulates while the conversion of carbon dioxide to plant biomass refers to the process of photosynthesis, taking CO2 and producing organic matter. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC25 -
AC26: 'Reply on RC8.17', Leigh-Ann Smit, 02 Feb 2026
Line 97- What type of biogeochemistry is referred to
This is referring to the chemical and biological processes that occur in the sediment that influences the decomposition of organic carbon. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC26 -
AC27: 'Reply on RC8.18', Leigh-Ann Smit, 02 Feb 2026
Line 99- Shouldn’t this be a different paragraph
In the manuscript it is already a different paragraph, but we will add subheadings to the introduction as stated in comment 8.3, with one being “Key carbon processes” which will help make the introduction clearer and more structured.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC27 -
AC28: 'Reply on RC8.19', Leigh-Ann Smit, 02 Feb 2026
Line 130- Change “marsh” to “marshes”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC28 -
AC29: 'Reply on RC8.20', Leigh-Ann Smit, 02 Feb 2026
Line 145- Why is there no subdivision of mangroves per continent?
We could not find literature/data on this, we could only source for salt marsh.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC29 -
AC30: 'Reply on RC8.21', Leigh-Ann Smit, 02 Feb 2026
Line 146- are salt marshes only storing a third SOC than mangroves?
Salt marshes do tend to have much lower organic carbon stocks per unit area than mangroves. We will double check the numbers to make sure they are correct. However, according to Reviewer 2 (comment 4.15), the paragraph from lines 134 to the end of the Table 1 starts to stray from the subject of this review, so we will be removing this section. This also allows for the introduction to be shorter and more streamlined.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC30 -
AC31: 'Reply on RC8.22', Leigh-Ann Smit, 02 Feb 2026
Line 186- This sentence is not universal or related to the previous one
We will remove the line “which includes burrow ventilation and particle reworking” to improve the flow of the paragraph.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC31 -
AC32: 'Reply on RC8.23', Leigh-Ann Smit, 02 Feb 2026
Line 196- Explain the difference between stabilizing and destabilizing
We will include a sentence to explain the difference.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC32 -
AC33: 'Reply on RC8.24', Leigh-Ann Smit, 02 Feb 2026
Line 198- Develop more this difference
We will include descriptions for upward conveyors, downward conveyors, biodiffusors and regenerators. This will include information on the feeding type, behaviour and life style of the organism which will explain how these characteristic determine the classification.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC33 -
AC34: 'Reply on RC8.25', Leigh-Ann Smit, 02 Feb 2026
Line 201- This sentence is not related to the others
The sentence “Collapsed burrows that are abandoned and become filled in, can be considered as indirect bioturbation (Kristensen et al., 2012).” will be relocated to line 186. After the sentence “Bioturbation involves any transport process performed by animals that affects sediment matrices, either directly or indirectly (Kristensen et al., 2012)”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC34 -
AC35: 'Reply on RC8.26', Leigh-Ann Smit, 02 Feb 2026
Line 215- Now you switch to crabs, more organization of the text is needed
We will restructure the text so that it is better organised and flows well, as several of the comments above and below help with.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC35 -
AC36: 'Reply on RC8.27', Leigh-Ann Smit, 02 Feb 2026
Line 240- Change “processes” to “process”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC36 -
AC37: 'Reply on RC8.28', Leigh-Ann Smit, 02 Feb 2026
Line 280- Does your statement apply to all salt marshes?
“In salt marshes, however, bioturbation had no significant effect on changes in surface elevation because they had lower levels of bioturbation compared to the mangroves.”. This is based on a study by Bennion et al. 2024 and their results do not apply to all salt marshes, no. We will clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC37 -
AC38: 'Reply on RC8.29', Leigh-Ann Smit, 02 Feb 2026
Line 294- Remove “Due to”
This will be removed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC38 -
AC39: 'Reply on RC8.30', Leigh-Ann Smit, 02 Feb 2026
Line 294- Replace “being” with “are”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC39 -
AC40: 'Reply on RC8.31', Leigh-Ann Smit, 02 Feb 2026
8.31) Line 294- Replace “they” to “and”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC40 -
AC41: 'Reply on RC8.32', Leigh-Ann Smit, 02 Feb 2026
Line 310- Why does a coarser sediment lead to more erosion?
Coarser sediment is generally non-cohesive so the sediment particles don’t stick together as well as what they would for fine sediment. We will add a sentence to the text to make it clear.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC41 -
AC42: 'Reply on RC8.33', Leigh-Ann Smit, 02 Feb 2026
8.33) Line 312- What benthic organisms?
This is referring specifically to bioturbating benthic species. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC42 -
AC43: 'Reply on RC8.34', Leigh-Ann Smit, 02 Feb 2026
Line 320- Why is roughness increasing erosion?
Roughness creates turbulence which can lead to increased erosion. We will adjust the text to make it clearer.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC43 -
AC44: 'Reply on RC8.35', Leigh-Ann Smit, 02 Feb 2026
Line 324- Clarify how Sesarma reticulatum causes increased erosion
Sesarma reticulatum denudes vegetation and produce burrows which facilitate erosional processes. This causes erosion and creek expansion. We will clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC44 -
AC45: 'Reply on RC8.36', Leigh-Ann Smit, 02 Feb 2026
Line 327- Separate chapter?
Although this paragraph touches on sediment stability it mainly speaks to sediment cohesion and erosion and we think fits well under this section “1.4.2 Erosion”. We will check the flow of all paragraphs and sections to improve coherency.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC45 -
AC46: 'Reply on RC8.37', Leigh-Ann Smit, 02 Feb 2026
Line 351- Sediment trapping should be looked at in a separate paragraph
We agree and will add a separate subheading which will focus on sediment trapping.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC46 -
AC47: 'Reply on RC8.38', Leigh-Ann Smit, 02 Feb 2026
Line 355- What is the difference between deposition and accretion
Deposition is the process of particles settling while accretion is the net vertical increase in surface elevation.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC47 -
AC48: 'Reply on RC8.39', Leigh-Ann Smit, 02 Feb 2026
Line 380- A table with all deposition measured in burrows would be fine here
We agree with this addition, and will add the table to this section.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC48 -
AC49: 'Reply on RC8.40', Leigh-Ann Smit, 02 Feb 2026
Line 414- Move “Sapelo Island” to the beginning of the sentence
We will change the sentence to “In Sapelo Island, Georgia, U.S it was estimated that Uca pugnax (a species of fiddler crab native to salt marshes along the coast of North America) excavated an amount of carbon that is equal to 20 % of what S. alterniflora produces belowground annually (Montague 1982; Table S1, Fig.3).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC49 -
AC50: 'Reply on RC8.41', Leigh-Ann Smit, 02 Feb 2026
Line 468- The effect of SLR might require a different chapter, how does it affect the other processes?
Sea level rise affects not just carbon but sediment processes too. It can influence erosion processes as well as chemical processes such as shifts in redox potential. Section 1.6 (Global change impacts on tidal marsh bioturbation) focuses more on this topic. We will move lines 468-472 to section 1.6.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC50 -
AC51: 'Reply on RC8.42', Leigh-Ann Smit, 02 Feb 2026
Line 487- This paragraph is not related to bioturbation
We will remove this paragraph (lines 487-493).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC51 -
AC52: 'Reply on RC8.43', Leigh-Ann Smit, 02 Feb 2026
Line 495- Remove “globally” and add “global” and change its position
We will edit our sentence and change it to “Mangroves are specifically vulnerable to changes in temperature and precipitation regimes, because the global distribution range is linked to sea surface temperature”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC52 -
AC53: 'Reply on RC8.44', Leigh-Ann Smit, 02 Feb 2026
Line 496- Replace “and this by” with “corresponding to”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC53 -
AC54: 'Reply on RC8.45', Leigh-Ann Smit, 02 Feb 2026
Line 508- Add the word “the” before Bahamas
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC54 -
AC55: 'Reply on RC8.46', Leigh-Ann Smit, 02 Feb 2026
Line 508- what is the link to bioturbation?
We did not link this paragraph (lines 494-508) very well to bioturbation. We will add information to make it clearer. We will link the climate driven shift in vegetation with bioturbator community shifts and the consequence on carbon storage.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC55 -
AC56: 'Reply on RC8.47', Leigh-Ann Smit, 02 Feb 2026
Line 515- Does this refer to only crabs or other bioturbators
No, changes in water levels and temperature can affect the behaviour and distribution of other bioturbators as well, not just crabs. We will adjust to text to make it clear and add examples of how other bioturbators are also influenced. For example, Zhou et al. 2022 found that thermal stress alters the burrowing depth and activity of the cockle Cerastoderma edule.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC56 -
AC57: 'Reply on RC8.48', Leigh-Ann Smit, 02 Feb 2026
Line 518- Make A in Alterniflora lowercase
We will correct this.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC57 -
AC58: 'Reply on RC8.49', Leigh-Ann Smit, 02 Feb 2026
Line 542- This paragraph is out of context, it does not relate to climate change
This is a whole new section, 1.7 (Synthesis and way forward). This paragraph is meant as a conclusion and synthesis of the whole manuscript, it is not specific to the previous section 1.6 (Global change impacts on tidal marsh bioturbation). We will ensure that this is clear in future drafts.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC58 -
AC59: 'Reply on RC8.50', Leigh-Ann Smit, 02 Feb 2026
Line 545- Change “influence” to “affect”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC59 -
AC60: 'Reply on RC8.51', Leigh-Ann Smit, 02 Feb 2026
Line 549- What species
This refers to species that produce an organic coating in the burrow walls, aiding with sediment cohesion, such as Sesarma reticulatum. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC60 -
AC61: 'Reply on RC8.52', Leigh-Ann Smit, 02 Feb 2026
Line 571- How?
This line “sediment-species interactions also have an influence on carbon cycling in tidal marshes” is referring to their ability to influence organic matter burial and decomposition rates. We will add this to the sentence to make it clearer.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC61 -
AC62: 'Reply on RC8.53', Leigh-Ann Smit, 02 Feb 2026
Line 578- Edit the sentence
We will change the sentence from “By advancing our understanding, management and restoration efforts could be improved, and better predict the resilience of tidal marshes under future climate change pressures” to “By advancing our understanding, we can improve management and restoration efforts and better predict the resilience of tidal wetlands under future climate change pressures” as suggested.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC62 -
AC63: 'Reply on RC8.54', Leigh-Ann Smit, 02 Feb 2026
Lines 45 and 243- Remove the comma
We will remove the commas from both sentences.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC63
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AC1: 'Reply on Reviewer 1's general comment', Leigh-Ann Smit, 02 Feb 2026
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RC2: 'Comment on egusphere-2025-4749', Anonymous Referee #2, 05 Jan 2026
This manuscript presents an overview of the impact of bioturbation on sediment and carbon sequestration impacts in tidal wetlands. The topic warrants increased attention, as the manuscript successfully shows, however the text requires some reorganization before publication.
- Sections should be more specific in some cases, and text within each section should be limited to that topic. For example, the introduction could be split into subsections of tidal marsh background, sediment processes, and carbon processes. Further, text organization within sections should follow a stricter pattern so the reader knows what to expect in each section. This is noted throughout the specific comments document.
- The text largely describes results of previous studies and would benefit from a stronger framework for comparison and synthesis (types of bioturbators, characteristics of wetlands, etc.). This might allow the text and the reader to draw stronger conclusions.
- The manuscript contains text in some sections that is not necessary and adds to the length. These instances are noted in the specific comments.
Specific comments and technical corrections: see attached file
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AC64: 'Reply on RC2's general comment', Leigh-Ann Smit, 02 Feb 2026
This manuscript presents an overview of the impact of bioturbation on sediment and carbon sequestration impacts in tidal wetlands. The topic warrants increased attention, as the manuscript successfully shows, however the text requires some reorganization before publication.
Thank you for this feedback and for the useful comments provided, which we have addressed below. In light of much of the feedback from Reviewer 1 as well, we have especially focussed on restructuring the text to ensure better organisation of the ideas presented.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC64 -
AC65: 'Reply on RC1', Leigh-Ann Smit, 02 Feb 2026
Sections should be more specific in some cases, and text within each section should be limited to that topic. For example, the introduction could be split into subsections of tidal marsh background, sediment processes, and carbon processes. Further, text organization within sections should follow a stricter pattern so the reader knows what to expect in each section. This is noted throughout the specific comments document.
Thank you for this constructive feedback. We will restructure and reorganize the manuscript by adding the suggested subheadings to the introduction section and ensuring each section focuses on one specific topic. We will relocate or remove text that does not align with a given topic/section throughout the manuscript. Please see our responses below to the comments provided in the reviewer’s annotated PDF.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC65 -
AC66: 'Reply on RC2', Leigh-Ann Smit, 02 Feb 2026
The text largely describes results of previous studies and would benefit from a stronger framework for comparison and synthesis (types of bioturbators, characteristics of wetlands, etc.). This might allow the text and the reader to draw stronger conclusions.
Based on feedback from reviewer 1, we will be adding two summary tables of bioregional bioturbators as well as bioturbators traits versus their influence on sediment and carbon dynamics which will help improve the framework as the reviewer suggests.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC66 -
AC67: 'Reply on RC3', Leigh-Ann Smit, 02 Feb 2026
The manuscript contains text in some sections that is not necessary and adds to the length. These instances are noted in the specific comments.
Thank you, we will review the comments in the document and remove or condense the text as much as possible, as described further below.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC67 -
AC68: 'Reply on RC4', Leigh-Ann Smit, 02 Feb 2026
Specific comments and technical corrections: see attached file
We note the very useful 43 edits and comments made by the reviewer throughout the PDF manuscript that assist us in addressing the three broad comments above, and have prepared the following responses to these:
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC68 -
AC69: 'Reply on RC4.1', Leigh-Ann Smit, 02 Feb 2026
Line 37- I suggest consistency between the terms tidal wetlands, salt marsh, and mangrove
We will go through the whole manuscript and ensure we remain consistent with these terms.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC69 -
AC70: 'Reply on RC4.2', Leigh-Ann Smit, 02 Feb 2026
Line 37- The introduction could be strengthened with some reworking. Currently, description of wetlands is long and needs to be tied to bioturbators. Additionally, some broad grouping of wetlands (macrotidal/microtidal, minerogenic/organogenic, marsh platform/creekbank) would help to later summarise impacts. This section could be split into 1) sediment/geomorphology and 2) carbon processes for more organisation
We agree that having section/subheadings in the introduction will help with the organisation. We will include subheadings such as “An overview of tidal wetlands”, “Key sediment processes” and “Key carbon processes”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC70 -
AC71: 'Reply on RC4.3', Leigh-Ann Smit, 02 Feb 2026
Line 52- Mcloud et al. 2011 would be a more appropriate citation, or move this citation to the end of the sentence
We will move the citation “Duarte De Paula Costa and Macreadie 2022” to the end of the sentence and replace its current position with Mcloud et al. 2011.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC71 -
AC72: 'Reply on RC4.4', Leigh-Ann Smit, 02 Feb 2026
Line 57- Are mudflats considered as a tidal marsh for this review? Above only salt marshes and mangroves are listed
Yes, mudflats are considered a tidal marsh for this review, however, the focus is more on salt marshes and mangroves given the historical focus on these habitats in terms of carbon processes, for example. However, we will include mudflats to the paragraph above and ensure that appropriate discussion is given to these habitats where relevant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC72 -
AC73: 'Reply on RC4.5', Leigh-Ann Smit, 02 Feb 2026
Line 67- Sentence structure here is awkward. Consider restructuring to place the important substance early I.e. “Large-scale change in tidal wetland area can be estimated with remote sensing data”
We will restructure the sentence from “By means of satellite observations, looking at changes in water presence, land loses, and gains can be estimated. It is estimated that 28 000 km2 of land has been eroded in tidal wetlands, which is double that of land gained” to “Large-scale change in tidal wetland area can be estimated with remote sensing data” as suggested.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC73 -
AC74: 'Reply on RC4.6', Leigh-Ann Smit, 02 Feb 2026
Line 70- There are more recent and wider scale accretion studies, such as https://doi.org/10.1007/s12237-014-9872-8, https://doi.org/10.1007/s12237-024-01332-z, or https://doi.org/10.1007/s12237-022-01141-2
We will remove the citation “Van Wijnen and Bakker, 2001” and replace it with more recent literature such as those suggested, thank you.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC74 -
AC75: 'Reply on RC4.7', Leigh-Ann Smit, 02 Feb 2026
Line- 76 Citation needed-there can be complex interactions, for example high water level can reduce erosion during a storm
We will include the reference Pang, T., Wang, X., Nawaz, R.A., Keefe, G. and Adekanmbi, T., 2023. Coastal erosion and climate change: A review on coastal-change process and modeling. Ambio, 52(12), pp.2034-2052 or Masselink, G. and Russell, P., 2013. Impacts of climate change on coastal erosion. MCCIP science review, 2013, pp.71-86. However, we acknowledge that there can be complex interactions and will acknowledge this in text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC75 -
AC76: 'Reply on RC4.8', Leigh-Ann Smit, 02 Feb 2026
Line 78- This paragraph is wordy and would be a candidate to cut back to shorten the introduction
We will be more concise and remove unnecessary information in this paragraph. For example, lines 93-96 can be removed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC76 -
AC77: 'Reply on RC4.9', Leigh-Ann Smit, 02 Feb 2026
Line 97- This and the following paragraph could be combined for more cohesive flow
We agree and will combine the two paragraphs for a better flow.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC77 -
AC78: 'Reply on RC4.10', Leigh-Ann Smit, 02 Feb 2026
Line 116- This should be compared to other vegetation types
We will add a sentence so that carbon stocks are compared between shrubby salt marsh and herbaceous salt marsh e.g. grasses and sedges.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC78 -
AC79: 'Reply on RC4.11', Leigh-Ann Smit, 02 Feb 2026
Line 119- Restoration is not mentioned elsewhere, so this point could be removed
We will remove the sentence “Carbon storage has been shown to be higher in mature salt marshes compared to restored or new salt marshes” as suggested.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC79 -
AC80: 'Reply on RC4.12', Leigh-Ann Smit, 02 Feb 2026
Line 120- Rather than comparing higher/lower carbon stocks, this paragraph would be more effective by distilling trends into a few broadly supported statements
We will shift the focus of the paragraph to general trends in carbon storage rather than comparisons of high versus low. For example, we will change the sentence “Marshes that experienced rapid relative sea level rise during the late Holocene have higher concentrations of soil carbon compared to those that were subject to long periods of sea level stability” to “Periods of rapid relative sea level rise during the late Holocene are associated with higher soil carbon concentrations compared to periods of sea level stability”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC80 -
AC81: 'Reply on RC4.13', Leigh-Ann Smit, 02 Feb 2026
Line 128- Lower redox potential typically indicates more anaerobic conditions
Yes, that is correct, we will remove “less anaerobic conditions” as it is redundant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC81 -
AC82: 'Reply on RC4.14', Leigh-Ann Smit, 02 Feb 2026
Line 129- This study is of arid wetlands and caution should be taken in applying this broadly to blue carbon ecosystems
Thank you, we will make sure to specify that the study is related to arid wetlands.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC82 -
AC83: 'Reply on RC4.15', Leigh-Ann Smit, 02 Feb 2026
Line 137-This section starts to stray from the subject of this review here, I’d suggest removing this paragraph
This paragraph was intended to provide a continent level overview of tidal marsh extent and organic carbon stocks. Given the length of the introduction and the need to streamline it, we will remove this section.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC83 -
AC84: 'Reply on RC4.16', Leigh-Ann Smit, 02 Feb 2026
Line 167- Figure 2 and Table 2 are largely repetitive. Figure 2 takes up a lot of space without presenting much information, particularly since the diagram is not to scale. I suggest removing figure 2
We will add the dominant regional bioturbators to figure 2 as suggested by Reviewer 1, which will add more information to the figure and make it more relevant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC84 -
AC85: 'Reply on RC4.17', Leigh-Ann Smit, 02 Feb 2026
Line 171- Where are the key words or words related to these? Please provide exact search parameters
The key words relating to tidal wetlands were: tidal wetland, salt marsh, mangrove,
The key words relating to carbon were: carbon stocks, carbon sequestration, carbon storage
The key words relating to sediment were: sediment dynamics, sediment accretion, sediment erosion, sedimentation.
We will add these key words to the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC85 -
AC86: 'Reply on RC4.18', Leigh-Ann Smit, 02 Feb 2026
Line 175- Where are the key words or words related to bioturbation
The key words relating to bioturbation were: bioturbation, bioturbator, soil animal, sediment mixing. We will add these key words to the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC86 -
AC87: 'Reply on RC4.19', Leigh-Ann Smit, 02 Feb 2026
Line 178- Over 10% of papers with seemingly broad search terms included bioturbation, this itself/alone doesn’t suggest to me that this is a poorly understood topic. This point would be strengthened if made after the bioturbation section, where the diversity of bioturbators is explained
We will move this sentence to section 1.3 (Bioturbation in tidal wetlands).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC87 -
AC88: 'Reply on RC4.20', Leigh-Ann Smit, 02 Feb 2026
Line 183- This paragraph has too many topics. I suggest the opening paragraph in this section be dedicated to defining bioturbation and identifying and classifying bioturbators. Impact of bioturbation can start in the next paragraph
Thank you, we will split the paragraph into two. The first paragraph will be dedicated to defining bioturbation and classifying bioturbators. The second paragraph will be for identifying the impact of bioturbation.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC88 -
AC89: 'Reply on RC4.21', Leigh-Ann Smit, 02 Feb 2026
Line 183- Vague, this could be strengthened by providing a count of studied bioturbators, or this sentence could be removed and the following sentence reformatted
Thank you, we will add a count of studied bioturbators to the sentence and if we cannot find a reliable number, we will remove the sentence and reformat the following sentence.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC89 -
AC90: 'Reply on RC4.22', Leigh-Ann Smit, 02 Feb 2026
Line 190- the definition of bioturbation should be at the start of the section
We will move the definition of bioturbation to the beginning of the paragraph.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC90 -
AC91: 'Reply on RC4.23', Leigh-Ann Smit, 02 Feb 2026
Line 195- Later in the text, burrow architecture becomes key context in differentiating impacts. Different architectures should be defined here
We will add a sentence explaining the different types of burrow architecture (e.g simple burrow, branched burrow, chambered burrow, multi entrance burrow).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC91 -
AC92: 'Reply on RC4.24', Leigh-Ann Smit, 02 Feb 2026
Line 201- What are destabilising and stabilising organisms? Some description should be given to these categories
Destabilizing organisms are those that disturb and mix sediment, making it looser and more erodible. Typically burrowing organisms. These include molluscs and crustaceans for example. Stabilising organisms are those that bind and compact sediment, increasing its resistance to erosion. These include organisms such as tube building polychaetes for example who produce mucus to bind the sediment. We will add a description to the text to give more context.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC92 -
AC93: 'Reply on RC4.25', Leigh-Ann Smit, 02 Feb 2026
Line 210- The outline of this sentence can help organize the following paragraphs. Physical, chemical, and biological impacts should each be described separately throughout each of the following sections
We will ensure these 3 aspects are covered in the sections that follow (accretion, erosion, sediment transport and deposition), and add information if it is not covered already.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC93 -
AC94: 'Reply on RC4.26', Leigh-Ann Smit, 03 Feb 2026
Line 212- This paragraph starts to get specific, and most of these points could be worked into the following sections, where this level of detail is more warranted
We will remove the paragraph and incorporate the relevant sentences into the following sections, where this level of detail is more appropriate.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC94 -
AC95: 'Reply on RC4.27', Leigh-Ann Smit, 03 Feb 2026
Line 241- Figure 3 has strong potential, and would benefit from stronger text support and some figure reorganization. Stronger text support: currently the text is organised using impacts of bioturbation, while figure 3 is organised by types of bioturbation and mechanisms that drive the impacts. This makes the figure difficult to follow along with through the text. Organising the figure in the same structure as the text would improve legibility. Figure reorganisation: the lower diagram doesn’t line up visual representations in the upper diagram, and this makes the figure difficult to follow. There are also some elements that are small/difficult to read in print. Also, the box for bioturbation impact #5 is blue, while the rest are grey- I don’t see an explanation for this in the caption
We will focus on incorporating Figure 3 more coherently in the text. Based on comment 2 from Reviewer 1, we will be adding a table which will include physical mechanisms of burrow construction and maintenance, looking at compaction, bioirrigation and sediment displacement. This will also include how these processes differ across different taxa and substrates. This will improve the link between the figure and the text. We will also reorganize the figure so that it follows the same structure as the text to ensure legibility. We will resize the elements in the figure so that they are legible. The blue box associated with #5 represents carbon and the grey shading represents sediment. We will include this explanation to the figure caption.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC95 -
AC96: 'Reply on RC4.28', Leigh-Ann Smit, 03 Feb 2026
Line 243- This is the opening paragraph of section on bioturbation, yet doesn’t mention bioturbation. This information should be moved, perhaps to a ‘tidal wetland geomorphology’ section in the introduction
We agree, we will relocate the first paragraph, lines 243-254, to the introduction section.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC96 -
AC97: 'Reply on RC4.29', Leigh-Ann Smit, 03 Feb 2026
Line 244- This is more of a focus for submerged vegetation such as seagrasses, which are not considered in this review. Regular inundation dramatically reduces salt marsh photosynthetic capacity
Thank you, we will remove that part of the sentence since it doesn’t make sense in the context of this review.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC97 -
AC98: 'Reply on RC4.30', Leigh-Ann Smit, 03 Feb 2026
Line 256- “extended periods and across surrounding areas” is vague
By “extended periods” we refer to long term effects, occurring over seasonal or multi-year timescale, not just over a single tidal cycle. By “surrounding areas” we refer to km-scale impacts, not just the immediate vicinity of the organism’s burrow. We will clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC98 -
AC99: 'Reply on RC4.31', Leigh-Ann Smit, 03 Feb 2026
Line 263- Are these bioturbators?
No, microphytobenthos (MPB) are not bioturbators. The inclusion of MPB in the text was meant as a comparison between some crab species that produce extracellular polymetric substances, such as mucus, which stabilises the sediment, in the same way that MPB stabilises the sediment. Later in the manuscript it is also mentioned that some bioturbators, such as crabs, consume MPB which then contributes to the destabilisation of sediment.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC99 -
AC100: 'Reply on RC4.32', Leigh-Ann Smit, 03 Feb 2026
Line 268- This doesn’t quite fit in the accretion section. Perhaps this is a better fit in the carbon sequestration, or in a soil chemistry section
The sentence “Low to moderate levels of bioturbation can be beneficial to primary productivity (Kristensen et al. 2008)” will be moved to the introduction section under the subheading Key carbon processes.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC100 -
AC101: 'Reply on RC4.33', Leigh-Ann Smit, 03 Feb 2026
Line 281- Slightly repetitive in this placement-this could be moved earlier
The line “the composition of crabs has the potential to influence ecosystems differently (Agusto et al. 2021)” will be moved to the paragraph above where it is currently positioned.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC101 -
AC102: 'Reply on RC4.34', Leigh-Ann Smit, 03 Feb 2026
Line 285- This statement doesn’t match the table and seems to be specific to this study-if so, that should be clarified
Yes, the statement “In salt marshes, however, bioturbation had no significant effect on changes in surface elevation” is specific to the study. We will ensure we clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC102 -
AC103: 'Reply on RC4.35', Leigh-Ann Smit, 03 Feb 2026
Line 322- is this always true? Earlier it was written that burrowers can enhance sediment cohesion
The statement “Burrowing activities weaken mud and clay banks in tidal marshes” is generally true. There are exceptions where some species e.g. Sesarma reticulatum secrete mucus which they coat their burrow walls with, which in turn can locally increase sediment cohesion. We will make this clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC103 -
AC104: 'Reply on RC4.36', Leigh-Ann Smit, 03 Feb 2026
Line 361- this is somewhat repetitive with sediment accretion and erosion above
We will review section 1.4.3 (sediment transport and deposition) to remove redundant content and make sure the information presented in the section provides additional insight.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC104 -
AC105: 'Reply on RC4.37', Leigh-Ann Smit, 03 Feb 2026
Line 415- Paragraph should have a more descriptive topic sentence
We will ensure the first sentence is more descriptive. We will alter it along the lines of “Consumers play a critical role in regulating carbon cycling, by directly and indirectly altering their physical environment, including primary production and sediment processes”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC105 -
AC106: 'Reply on RC4.38', Leigh-Ann Smit, 03 Feb 2026
Line 441- Could this also aerate and increase decomposition of previously stable C?
Yes, it could, we will include this information to that sentence, referencing Wang et al. 2010, who showed that bioturbation by crabs can promote sediment and carbon turnover (Wang, J. Q., Zhang, X. D., Jiang, L. F., Bertness, M. D., Fang, C. M., Chen, J. K., Hara, T., and Li, B.: Bioturbation of Burrowing Crabs Promotes Sediment Turnover and Carbon and Nitrogen Movements in an Estuarine Salt Marsh, Ecosystems, 13, 586– 599, https://doi.org/10.1007/s10021-010-9342-5, 2010.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC106 -
AC107: 'Reply on RC4.39', Leigh-Ann Smit, 03 Feb 2026
Line 488- This can also be from biological drivers
We agree that surface elevation can also increase as a result of biological drivers. We will edit the text to make it clear that both physical and biological processes contribute.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC107 -
AC108: 'Reply on RC4.40', Leigh-Ann Smit, 03 Feb 2026
Line 490- Repetitive with above
We will remove lines 490-495 as they are redundant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC108 -
AC109: 'Reply on RC4.41', Leigh-Ann Smit, 03 Feb 2026
Line 502- Winter freeze occurrence
We will include winter freeze occurrence to the sentence, it will read “Mangroves are specifically vulnerable to changes in temperature and precipitation regimes, because the distribution range globally is linked to sea surface temperature and the occurrence of winter freezes”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC109 -
AC110: 'Reply on RC4.42', Leigh-Ann Smit, 03 Feb 2026
Line 504- The reference is missing
Thank you, the reference is: Tomlinson, P.B., 1999. The Botany of Mangroves. Cambridge University Press, Cambridge, United Kingdom. We will be sure to add it to the reference list.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC110 -
AC111: 'Reply on RC4.43', Leigh-Ann Smit, 03 Feb 2026
Line 539- “Increases in drainage density is necessary to manage the expanding tidal prism and effectively drain the marsh surface to prevent waterlogging”- Abrupt shift in topic
We will improve the link between the sentence “Increases in drainage density is necessary to manage the expanding tidal prism and effectively drain the marsh surface to prevent waterlogging” and crabs. The sentence will change to “Crab activity at tidal creeks can effectively increase drainage density, helping to manage the expanding tidal prism and prevent waterlogging on the marsh surface (Farron et al. 2020)”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC111
Status: closed
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RC1: 'Comment on egusphere-2025-4749', Anonymous Referee #1, 12 Nov 2025
This manuscript presents an overview of the effect of bioturbation on tidal wetlands. While the topic is of great interest, the material is poorly organized, making it difficult for readers to develop a general understanding of the subject. As it stands, the manuscript reads more like a collection of disparate results from the literature rather than a cohesive synthesis. Below, I outline major points for improvement, along with specific suggested edits.
Major Points for Improvement:
- An overview of different bioturbators across continents and regions should be provided early in the paper.
- A detailed description of the physics of burrowing, and how it varies depending on species, substrate, and region, is currently missing and should be included.
- The introduction is fragmented and contains much extraneous information; it needs to be more focused on the central topic.
- Only two figures are presented, which is insufficient. The manuscript would benefit from additional conceptual diagrams illustrating main processes and feedbacks. Figures that summarize the text would be particularly helpful.
- Including tables that compile data from multiple studies to generalize results would greatly strengthen the manuscript.
- There is little attempt to synthesize the presented material. Synthesis should be provided throughout, not only in the conclusion, and could be effectively conveyed via summary plots or tables.
- A dedicated section or paragraph on species migration driven by global warming is missing and should be incorporated.
Some additional minor edits are noted in the text.
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AC1: 'Reply on Reviewer 1's general comment', Leigh-Ann Smit, 02 Feb 2026
This manuscript presents an overview of the effect of bioturbation on tidal wetlands. While the topic is of great interest, the material is poorly organized, making it difficult for readers to develop a general understanding of the subject. As it stands, the manuscript reads more like a collection of disparate results from the literature rather than a cohesive synthesis. Below, I outline major points for improvement, along with specific suggested edits.
Thank you for this feedback and for the useful comments provided, which we have addressed below. We will especially focus on restructuring the text to ensure flow and coherency.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC1 -
AC2: 'Reply on RC1', Leigh-Ann Smit, 02 Feb 2026
An overview of different bioturbators across continents and regions should be provided early in the paper.
We agree that adding this information early on in the paper would improve the contextual framing of the review. We will add this to section 1.3 (Bioturbation in coastal wetlands). This section introduces the organisms associated with bioturbation, so we will incorporate the types of bioturbators and where they are found. This will highlight the typical organisms (down to species-level where information is available) found across the tidal wetlands of global bioregions. A new table will be included to summarise the information.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC2 -
AC3: 'Reply on RC2', Leigh-Ann Smit, 02 Feb 2026
A detailed description of the physics of burrowing, and how it varies depending on species, substrate, and region, is currently missing and should be included.
We will expand the manuscript to provide this information as suggested. We will include physical mechanisms of burrow construction and maintenance, looking at compaction, bioirrigation and sediment displacement. This will also include how these processes differ across different taxa and substrates. This will be added to section 1.3 (Bioturbation in coastal tidal wetlands). A new table will be included to summarise this information.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC3 -
AC4: 'Reply on RC3', Leigh-Ann Smit, 02 Feb 2026
The introduction is fragmented and contains much extraneous information; it needs to be more focused on the central topic.
We will reorganize and streamline the introduction by adding subheadings and removing unnecessary information. We will include headings such as “An overview of tidal wetlands”, “Key sediment processes” and “Key carbon processes”, as suggested by reviewer 2 (comment 2.1).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC4 -
AC5: 'Reply on RC4', Leigh-Ann Smit, 02 Feb 2026
Only two figures are presented, which is insufficient. The manuscript would benefit from additional conceptual diagrams illustrating main processes and feedbacks. Figures that summarize the text would be particularly helpful.
Although Figure 3 summarises many of the processes influenced by bioturbators, we will add additional illustrations to improve clarity and highlight key processes, especially in line with comment 1.2 above to summarise these. We will add the dominant regional bioturbators to Figure 2 and create an additional figure on the burrowing modes and effects on sediment of different bioturbators (e.g. crab burrows, worm burrows, shrimp burrows). We will also include a conceptual summary figure at the end of the manuscript with the major feedbacks, both positive and negative and link it to sediment dynamics and carbon processes.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC5 -
AC6: 'Reply on RC5', Leigh-Ann Smit, 02 Feb 2026
Including tables that compile data from multiple studies to generalize results would greatly strengthen the manuscript.
Table S1 provides a synthesis of the literature and studies provided in the text. We will draft two summary data tables in response to comments 1.1 and 1.2 above, which will also thereby summarise the information presented in Table S1.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC6 -
AC7: 'Reply on RC6', Leigh-Ann Smit, 02 Feb 2026
Including tables that compile data from multiple studies to generalize results would greatly strengthen the manuscript.
Table S1 provides a synthesis of the literature and studies provided in the text. We will draft two summary data tables in response to comments 1.1 and 1.2 above, which will also thereby summarise the information presented in Table S1.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC7 -
AC8: 'Reply on RC7', Leigh-Ann Smit, 02 Feb 2026
A dedicated section or paragraph on species migration driven by global warming is missing and should be incorporated.
We will add an additional paragraph to section 1.6 (Global change impacts on tidal marsh bioturbation), to include this information. We will focus on projected shifts in the distribution of species and how this might influence bioturbation patterns, with implications for sediment and carbon processes under these scenarios.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC8 -
AC9: 'Reply on RC7', Leigh-Ann Smit, 02 Feb 2026
Some additional minor edits are noted in the text.
We note the very useful 54 edits and comments made by this reviewer throughout the PDF manuscript that assist us in addressing the seven broad comments above, and have prepared the following responses to these:
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC9 -
AC10: 'Reply on RC8.1', Leigh-Ann Smit, 02 Feb 2026
Line 2-Replace “marshes” with “wetlands”.
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC10 -
AC11: 'Reply on RC8.2', Leigh-Ann Smit, 02 Feb 2026
Line 17- Replace “influence” with “affect”.
This will be changed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC11 -
AC12: 'Reply on RC8.3', Leigh-Ann Smit, 02 Feb 2026
Line 37- The introduction is too fragmented with too much information, you might want to reduce it and make it more focused
We will reorganise the text and include headings such as “An overview of tidal wetlands”, “Key sediment processes” and “Key carbon processes”. We will also remove any unnecessary information.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC12 -
AC13: 'Reply on RC8.4', Leigh-Ann Smit, 02 Feb 2026
Line 39- Replace “marshes” with “wetlands”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC13 -
AC14: 'Reply on RC8.5', Leigh-Ann Smit, 02 Feb 2026
Line 40- Change “exist between” to “connect”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC14 -
AC15: 'Reply on RC8.6', Leigh-Ann Smit, 02 Feb 2026
Line 43- Replace “they” with “Tidal wetlands”
This will be changed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC15 -
AC16: 'Reply on RC8.7', Leigh-Ann Smit, 02 Feb 2026
Line 50- Replace “with” with “and”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC16 -
AC17: 'Reply on RC8.8', Leigh-Ann Smit, 02 Feb 2026
Line 55- Replace “their” with “its”
This will be replaced.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC17 -
AC18: 'Reply on RC8.9', Leigh-Ann Smit, 02 Feb 2026
Line 57- Include a space between the words “abiotic” and “and”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC18 -
AC19: 'Reply on RC8.10', Leigh-Ann Smit, 02 Feb 2026
Line 57- Remove “s” at the end of includes
We will change it to “include”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC19 -
AC20: 'Reply on RC8.11', Leigh-Ann Smit, 02 Feb 2026
Line 62- Replace “loses” with “losses”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC20 -
AC21: 'Reply on RC8.12', Leigh-Ann Smit, 02 Feb 2026
Line 62- Change “estimated” to “assessed”
This will be changed
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC21 -
AC22: 'Reply on RC8.13', Leigh-Ann Smit, 02 Feb 2026
Line 62- Reference needed
The reference “Mentaschi et al. 2018” will be added.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC22 -
AC23: 'Reply on RC8.14', Leigh-Ann Smit, 02 Feb 2026
Line 63- This paragraph “Some studies have revealed that accretion rates are insufficient for tidal marshes to keep pace with sea level rise (e.g. Van Wijnen and Bakker, 2001), while others have found that accretion rates are high enough to keep pace with moderate rises in sea level (e.g. Morris et al., 2002)” is out of context
This section will be moved to section 1.6 (Global change impacts on tidal marsh bioturbation).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC23 -
AC24: 'Reply on RC8.15', Leigh-Ann Smit, 02 Feb 2026
Line 82- Provide clarity on what is meant by relative to the ocean carbon cycle
This will be made clearer in the text as follows “These blue carbon habitats store up to 70% of carbon, relative to the ocean carbon cycle, despite covering less than 2% of the area of the global ocean”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC24 -
AC25: 'Reply on RC8.16', Leigh-Ann Smit, 02 Feb 2026
Line 96- What is the difference between high productivity and conversion of carbon dioxide into plant biomass
High productivity is referring to how much biomass accumulates while the conversion of carbon dioxide to plant biomass refers to the process of photosynthesis, taking CO2 and producing organic matter. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC25 -
AC26: 'Reply on RC8.17', Leigh-Ann Smit, 02 Feb 2026
Line 97- What type of biogeochemistry is referred to
This is referring to the chemical and biological processes that occur in the sediment that influences the decomposition of organic carbon. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC26 -
AC27: 'Reply on RC8.18', Leigh-Ann Smit, 02 Feb 2026
Line 99- Shouldn’t this be a different paragraph
In the manuscript it is already a different paragraph, but we will add subheadings to the introduction as stated in comment 8.3, with one being “Key carbon processes” which will help make the introduction clearer and more structured.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC27 -
AC28: 'Reply on RC8.19', Leigh-Ann Smit, 02 Feb 2026
Line 130- Change “marsh” to “marshes”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC28 -
AC29: 'Reply on RC8.20', Leigh-Ann Smit, 02 Feb 2026
Line 145- Why is there no subdivision of mangroves per continent?
We could not find literature/data on this, we could only source for salt marsh.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC29 -
AC30: 'Reply on RC8.21', Leigh-Ann Smit, 02 Feb 2026
Line 146- are salt marshes only storing a third SOC than mangroves?
Salt marshes do tend to have much lower organic carbon stocks per unit area than mangroves. We will double check the numbers to make sure they are correct. However, according to Reviewer 2 (comment 4.15), the paragraph from lines 134 to the end of the Table 1 starts to stray from the subject of this review, so we will be removing this section. This also allows for the introduction to be shorter and more streamlined.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC30 -
AC31: 'Reply on RC8.22', Leigh-Ann Smit, 02 Feb 2026
Line 186- This sentence is not universal or related to the previous one
We will remove the line “which includes burrow ventilation and particle reworking” to improve the flow of the paragraph.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC31 -
AC32: 'Reply on RC8.23', Leigh-Ann Smit, 02 Feb 2026
Line 196- Explain the difference between stabilizing and destabilizing
We will include a sentence to explain the difference.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC32 -
AC33: 'Reply on RC8.24', Leigh-Ann Smit, 02 Feb 2026
Line 198- Develop more this difference
We will include descriptions for upward conveyors, downward conveyors, biodiffusors and regenerators. This will include information on the feeding type, behaviour and life style of the organism which will explain how these characteristic determine the classification.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC33 -
AC34: 'Reply on RC8.25', Leigh-Ann Smit, 02 Feb 2026
Line 201- This sentence is not related to the others
The sentence “Collapsed burrows that are abandoned and become filled in, can be considered as indirect bioturbation (Kristensen et al., 2012).” will be relocated to line 186. After the sentence “Bioturbation involves any transport process performed by animals that affects sediment matrices, either directly or indirectly (Kristensen et al., 2012)”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC34 -
AC35: 'Reply on RC8.26', Leigh-Ann Smit, 02 Feb 2026
Line 215- Now you switch to crabs, more organization of the text is needed
We will restructure the text so that it is better organised and flows well, as several of the comments above and below help with.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC35 -
AC36: 'Reply on RC8.27', Leigh-Ann Smit, 02 Feb 2026
Line 240- Change “processes” to “process”
This will be corrected.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC36 -
AC37: 'Reply on RC8.28', Leigh-Ann Smit, 02 Feb 2026
Line 280- Does your statement apply to all salt marshes?
“In salt marshes, however, bioturbation had no significant effect on changes in surface elevation because they had lower levels of bioturbation compared to the mangroves.”. This is based on a study by Bennion et al. 2024 and their results do not apply to all salt marshes, no. We will clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC37 -
AC38: 'Reply on RC8.29', Leigh-Ann Smit, 02 Feb 2026
Line 294- Remove “Due to”
This will be removed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC38 -
AC39: 'Reply on RC8.30', Leigh-Ann Smit, 02 Feb 2026
Line 294- Replace “being” with “are”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC39 -
AC40: 'Reply on RC8.31', Leigh-Ann Smit, 02 Feb 2026
8.31) Line 294- Replace “they” to “and”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC40 -
AC41: 'Reply on RC8.32', Leigh-Ann Smit, 02 Feb 2026
Line 310- Why does a coarser sediment lead to more erosion?
Coarser sediment is generally non-cohesive so the sediment particles don’t stick together as well as what they would for fine sediment. We will add a sentence to the text to make it clear.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC41 -
AC42: 'Reply on RC8.33', Leigh-Ann Smit, 02 Feb 2026
8.33) Line 312- What benthic organisms?
This is referring specifically to bioturbating benthic species. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC42 -
AC43: 'Reply on RC8.34', Leigh-Ann Smit, 02 Feb 2026
Line 320- Why is roughness increasing erosion?
Roughness creates turbulence which can lead to increased erosion. We will adjust the text to make it clearer.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC43 -
AC44: 'Reply on RC8.35', Leigh-Ann Smit, 02 Feb 2026
Line 324- Clarify how Sesarma reticulatum causes increased erosion
Sesarma reticulatum denudes vegetation and produce burrows which facilitate erosional processes. This causes erosion and creek expansion. We will clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC44 -
AC45: 'Reply on RC8.36', Leigh-Ann Smit, 02 Feb 2026
Line 327- Separate chapter?
Although this paragraph touches on sediment stability it mainly speaks to sediment cohesion and erosion and we think fits well under this section “1.4.2 Erosion”. We will check the flow of all paragraphs and sections to improve coherency.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC45 -
AC46: 'Reply on RC8.37', Leigh-Ann Smit, 02 Feb 2026
Line 351- Sediment trapping should be looked at in a separate paragraph
We agree and will add a separate subheading which will focus on sediment trapping.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC46 -
AC47: 'Reply on RC8.38', Leigh-Ann Smit, 02 Feb 2026
Line 355- What is the difference between deposition and accretion
Deposition is the process of particles settling while accretion is the net vertical increase in surface elevation.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC47 -
AC48: 'Reply on RC8.39', Leigh-Ann Smit, 02 Feb 2026
Line 380- A table with all deposition measured in burrows would be fine here
We agree with this addition, and will add the table to this section.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC48 -
AC49: 'Reply on RC8.40', Leigh-Ann Smit, 02 Feb 2026
Line 414- Move “Sapelo Island” to the beginning of the sentence
We will change the sentence to “In Sapelo Island, Georgia, U.S it was estimated that Uca pugnax (a species of fiddler crab native to salt marshes along the coast of North America) excavated an amount of carbon that is equal to 20 % of what S. alterniflora produces belowground annually (Montague 1982; Table S1, Fig.3).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC49 -
AC50: 'Reply on RC8.41', Leigh-Ann Smit, 02 Feb 2026
Line 468- The effect of SLR might require a different chapter, how does it affect the other processes?
Sea level rise affects not just carbon but sediment processes too. It can influence erosion processes as well as chemical processes such as shifts in redox potential. Section 1.6 (Global change impacts on tidal marsh bioturbation) focuses more on this topic. We will move lines 468-472 to section 1.6.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC50 -
AC51: 'Reply on RC8.42', Leigh-Ann Smit, 02 Feb 2026
Line 487- This paragraph is not related to bioturbation
We will remove this paragraph (lines 487-493).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC51 -
AC52: 'Reply on RC8.43', Leigh-Ann Smit, 02 Feb 2026
Line 495- Remove “globally” and add “global” and change its position
We will edit our sentence and change it to “Mangroves are specifically vulnerable to changes in temperature and precipitation regimes, because the global distribution range is linked to sea surface temperature”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC52 -
AC53: 'Reply on RC8.44', Leigh-Ann Smit, 02 Feb 2026
Line 496- Replace “and this by” with “corresponding to”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC53 -
AC54: 'Reply on RC8.45', Leigh-Ann Smit, 02 Feb 2026
Line 508- Add the word “the” before Bahamas
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC54 -
AC55: 'Reply on RC8.46', Leigh-Ann Smit, 02 Feb 2026
Line 508- what is the link to bioturbation?
We did not link this paragraph (lines 494-508) very well to bioturbation. We will add information to make it clearer. We will link the climate driven shift in vegetation with bioturbator community shifts and the consequence on carbon storage.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC55 -
AC56: 'Reply on RC8.47', Leigh-Ann Smit, 02 Feb 2026
Line 515- Does this refer to only crabs or other bioturbators
No, changes in water levels and temperature can affect the behaviour and distribution of other bioturbators as well, not just crabs. We will adjust to text to make it clear and add examples of how other bioturbators are also influenced. For example, Zhou et al. 2022 found that thermal stress alters the burrowing depth and activity of the cockle Cerastoderma edule.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC56 -
AC57: 'Reply on RC8.48', Leigh-Ann Smit, 02 Feb 2026
Line 518- Make A in Alterniflora lowercase
We will correct this.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC57 -
AC58: 'Reply on RC8.49', Leigh-Ann Smit, 02 Feb 2026
Line 542- This paragraph is out of context, it does not relate to climate change
This is a whole new section, 1.7 (Synthesis and way forward). This paragraph is meant as a conclusion and synthesis of the whole manuscript, it is not specific to the previous section 1.6 (Global change impacts on tidal marsh bioturbation). We will ensure that this is clear in future drafts.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC58 -
AC59: 'Reply on RC8.50', Leigh-Ann Smit, 02 Feb 2026
Line 545- Change “influence” to “affect”
Will do.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC59 -
AC60: 'Reply on RC8.51', Leigh-Ann Smit, 02 Feb 2026
Line 549- What species
This refers to species that produce an organic coating in the burrow walls, aiding with sediment cohesion, such as Sesarma reticulatum. This will be made clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC60 -
AC61: 'Reply on RC8.52', Leigh-Ann Smit, 02 Feb 2026
Line 571- How?
This line “sediment-species interactions also have an influence on carbon cycling in tidal marshes” is referring to their ability to influence organic matter burial and decomposition rates. We will add this to the sentence to make it clearer.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC61 -
AC62: 'Reply on RC8.53', Leigh-Ann Smit, 02 Feb 2026
Line 578- Edit the sentence
We will change the sentence from “By advancing our understanding, management and restoration efforts could be improved, and better predict the resilience of tidal marshes under future climate change pressures” to “By advancing our understanding, we can improve management and restoration efforts and better predict the resilience of tidal wetlands under future climate change pressures” as suggested.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC62 -
AC63: 'Reply on RC8.54', Leigh-Ann Smit, 02 Feb 2026
Lines 45 and 243- Remove the comma
We will remove the commas from both sentences.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC63
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RC2: 'Comment on egusphere-2025-4749', Anonymous Referee #2, 05 Jan 2026
This manuscript presents an overview of the impact of bioturbation on sediment and carbon sequestration impacts in tidal wetlands. The topic warrants increased attention, as the manuscript successfully shows, however the text requires some reorganization before publication.
- Sections should be more specific in some cases, and text within each section should be limited to that topic. For example, the introduction could be split into subsections of tidal marsh background, sediment processes, and carbon processes. Further, text organization within sections should follow a stricter pattern so the reader knows what to expect in each section. This is noted throughout the specific comments document.
- The text largely describes results of previous studies and would benefit from a stronger framework for comparison and synthesis (types of bioturbators, characteristics of wetlands, etc.). This might allow the text and the reader to draw stronger conclusions.
- The manuscript contains text in some sections that is not necessary and adds to the length. These instances are noted in the specific comments.
Specific comments and technical corrections: see attached file
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AC64: 'Reply on RC2's general comment', Leigh-Ann Smit, 02 Feb 2026
This manuscript presents an overview of the impact of bioturbation on sediment and carbon sequestration impacts in tidal wetlands. The topic warrants increased attention, as the manuscript successfully shows, however the text requires some reorganization before publication.
Thank you for this feedback and for the useful comments provided, which we have addressed below. In light of much of the feedback from Reviewer 1 as well, we have especially focussed on restructuring the text to ensure better organisation of the ideas presented.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC64 -
AC65: 'Reply on RC1', Leigh-Ann Smit, 02 Feb 2026
Sections should be more specific in some cases, and text within each section should be limited to that topic. For example, the introduction could be split into subsections of tidal marsh background, sediment processes, and carbon processes. Further, text organization within sections should follow a stricter pattern so the reader knows what to expect in each section. This is noted throughout the specific comments document.
Thank you for this constructive feedback. We will restructure and reorganize the manuscript by adding the suggested subheadings to the introduction section and ensuring each section focuses on one specific topic. We will relocate or remove text that does not align with a given topic/section throughout the manuscript. Please see our responses below to the comments provided in the reviewer’s annotated PDF.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC65 -
AC66: 'Reply on RC2', Leigh-Ann Smit, 02 Feb 2026
The text largely describes results of previous studies and would benefit from a stronger framework for comparison and synthesis (types of bioturbators, characteristics of wetlands, etc.). This might allow the text and the reader to draw stronger conclusions.
Based on feedback from reviewer 1, we will be adding two summary tables of bioregional bioturbators as well as bioturbators traits versus their influence on sediment and carbon dynamics which will help improve the framework as the reviewer suggests.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC66 -
AC67: 'Reply on RC3', Leigh-Ann Smit, 02 Feb 2026
The manuscript contains text in some sections that is not necessary and adds to the length. These instances are noted in the specific comments.
Thank you, we will review the comments in the document and remove or condense the text as much as possible, as described further below.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC67 -
AC68: 'Reply on RC4', Leigh-Ann Smit, 02 Feb 2026
Specific comments and technical corrections: see attached file
We note the very useful 43 edits and comments made by the reviewer throughout the PDF manuscript that assist us in addressing the three broad comments above, and have prepared the following responses to these:
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC68 -
AC69: 'Reply on RC4.1', Leigh-Ann Smit, 02 Feb 2026
Line 37- I suggest consistency between the terms tidal wetlands, salt marsh, and mangrove
We will go through the whole manuscript and ensure we remain consistent with these terms.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC69 -
AC70: 'Reply on RC4.2', Leigh-Ann Smit, 02 Feb 2026
Line 37- The introduction could be strengthened with some reworking. Currently, description of wetlands is long and needs to be tied to bioturbators. Additionally, some broad grouping of wetlands (macrotidal/microtidal, minerogenic/organogenic, marsh platform/creekbank) would help to later summarise impacts. This section could be split into 1) sediment/geomorphology and 2) carbon processes for more organisation
We agree that having section/subheadings in the introduction will help with the organisation. We will include subheadings such as “An overview of tidal wetlands”, “Key sediment processes” and “Key carbon processes”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC70 -
AC71: 'Reply on RC4.3', Leigh-Ann Smit, 02 Feb 2026
Line 52- Mcloud et al. 2011 would be a more appropriate citation, or move this citation to the end of the sentence
We will move the citation “Duarte De Paula Costa and Macreadie 2022” to the end of the sentence and replace its current position with Mcloud et al. 2011.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC71 -
AC72: 'Reply on RC4.4', Leigh-Ann Smit, 02 Feb 2026
Line 57- Are mudflats considered as a tidal marsh for this review? Above only salt marshes and mangroves are listed
Yes, mudflats are considered a tidal marsh for this review, however, the focus is more on salt marshes and mangroves given the historical focus on these habitats in terms of carbon processes, for example. However, we will include mudflats to the paragraph above and ensure that appropriate discussion is given to these habitats where relevant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC72 -
AC73: 'Reply on RC4.5', Leigh-Ann Smit, 02 Feb 2026
Line 67- Sentence structure here is awkward. Consider restructuring to place the important substance early I.e. “Large-scale change in tidal wetland area can be estimated with remote sensing data”
We will restructure the sentence from “By means of satellite observations, looking at changes in water presence, land loses, and gains can be estimated. It is estimated that 28 000 km2 of land has been eroded in tidal wetlands, which is double that of land gained” to “Large-scale change in tidal wetland area can be estimated with remote sensing data” as suggested.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC73 -
AC74: 'Reply on RC4.6', Leigh-Ann Smit, 02 Feb 2026
Line 70- There are more recent and wider scale accretion studies, such as https://doi.org/10.1007/s12237-014-9872-8, https://doi.org/10.1007/s12237-024-01332-z, or https://doi.org/10.1007/s12237-022-01141-2
We will remove the citation “Van Wijnen and Bakker, 2001” and replace it with more recent literature such as those suggested, thank you.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC74 -
AC75: 'Reply on RC4.7', Leigh-Ann Smit, 02 Feb 2026
Line- 76 Citation needed-there can be complex interactions, for example high water level can reduce erosion during a storm
We will include the reference Pang, T., Wang, X., Nawaz, R.A., Keefe, G. and Adekanmbi, T., 2023. Coastal erosion and climate change: A review on coastal-change process and modeling. Ambio, 52(12), pp.2034-2052 or Masselink, G. and Russell, P., 2013. Impacts of climate change on coastal erosion. MCCIP science review, 2013, pp.71-86. However, we acknowledge that there can be complex interactions and will acknowledge this in text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC75 -
AC76: 'Reply on RC4.8', Leigh-Ann Smit, 02 Feb 2026
Line 78- This paragraph is wordy and would be a candidate to cut back to shorten the introduction
We will be more concise and remove unnecessary information in this paragraph. For example, lines 93-96 can be removed.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC76 -
AC77: 'Reply on RC4.9', Leigh-Ann Smit, 02 Feb 2026
Line 97- This and the following paragraph could be combined for more cohesive flow
We agree and will combine the two paragraphs for a better flow.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC77 -
AC78: 'Reply on RC4.10', Leigh-Ann Smit, 02 Feb 2026
Line 116- This should be compared to other vegetation types
We will add a sentence so that carbon stocks are compared between shrubby salt marsh and herbaceous salt marsh e.g. grasses and sedges.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC78 -
AC79: 'Reply on RC4.11', Leigh-Ann Smit, 02 Feb 2026
Line 119- Restoration is not mentioned elsewhere, so this point could be removed
We will remove the sentence “Carbon storage has been shown to be higher in mature salt marshes compared to restored or new salt marshes” as suggested.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC79 -
AC80: 'Reply on RC4.12', Leigh-Ann Smit, 02 Feb 2026
Line 120- Rather than comparing higher/lower carbon stocks, this paragraph would be more effective by distilling trends into a few broadly supported statements
We will shift the focus of the paragraph to general trends in carbon storage rather than comparisons of high versus low. For example, we will change the sentence “Marshes that experienced rapid relative sea level rise during the late Holocene have higher concentrations of soil carbon compared to those that were subject to long periods of sea level stability” to “Periods of rapid relative sea level rise during the late Holocene are associated with higher soil carbon concentrations compared to periods of sea level stability”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC80 -
AC81: 'Reply on RC4.13', Leigh-Ann Smit, 02 Feb 2026
Line 128- Lower redox potential typically indicates more anaerobic conditions
Yes, that is correct, we will remove “less anaerobic conditions” as it is redundant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC81 -
AC82: 'Reply on RC4.14', Leigh-Ann Smit, 02 Feb 2026
Line 129- This study is of arid wetlands and caution should be taken in applying this broadly to blue carbon ecosystems
Thank you, we will make sure to specify that the study is related to arid wetlands.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC82 -
AC83: 'Reply on RC4.15', Leigh-Ann Smit, 02 Feb 2026
Line 137-This section starts to stray from the subject of this review here, I’d suggest removing this paragraph
This paragraph was intended to provide a continent level overview of tidal marsh extent and organic carbon stocks. Given the length of the introduction and the need to streamline it, we will remove this section.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC83 -
AC84: 'Reply on RC4.16', Leigh-Ann Smit, 02 Feb 2026
Line 167- Figure 2 and Table 2 are largely repetitive. Figure 2 takes up a lot of space without presenting much information, particularly since the diagram is not to scale. I suggest removing figure 2
We will add the dominant regional bioturbators to figure 2 as suggested by Reviewer 1, which will add more information to the figure and make it more relevant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC84 -
AC85: 'Reply on RC4.17', Leigh-Ann Smit, 02 Feb 2026
Line 171- Where are the key words or words related to these? Please provide exact search parameters
The key words relating to tidal wetlands were: tidal wetland, salt marsh, mangrove,
The key words relating to carbon were: carbon stocks, carbon sequestration, carbon storage
The key words relating to sediment were: sediment dynamics, sediment accretion, sediment erosion, sedimentation.
We will add these key words to the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC85 -
AC86: 'Reply on RC4.18', Leigh-Ann Smit, 02 Feb 2026
Line 175- Where are the key words or words related to bioturbation
The key words relating to bioturbation were: bioturbation, bioturbator, soil animal, sediment mixing. We will add these key words to the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC86 -
AC87: 'Reply on RC4.19', Leigh-Ann Smit, 02 Feb 2026
Line 178- Over 10% of papers with seemingly broad search terms included bioturbation, this itself/alone doesn’t suggest to me that this is a poorly understood topic. This point would be strengthened if made after the bioturbation section, where the diversity of bioturbators is explained
We will move this sentence to section 1.3 (Bioturbation in tidal wetlands).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC87 -
AC88: 'Reply on RC4.20', Leigh-Ann Smit, 02 Feb 2026
Line 183- This paragraph has too many topics. I suggest the opening paragraph in this section be dedicated to defining bioturbation and identifying and classifying bioturbators. Impact of bioturbation can start in the next paragraph
Thank you, we will split the paragraph into two. The first paragraph will be dedicated to defining bioturbation and classifying bioturbators. The second paragraph will be for identifying the impact of bioturbation.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC88 -
AC89: 'Reply on RC4.21', Leigh-Ann Smit, 02 Feb 2026
Line 183- Vague, this could be strengthened by providing a count of studied bioturbators, or this sentence could be removed and the following sentence reformatted
Thank you, we will add a count of studied bioturbators to the sentence and if we cannot find a reliable number, we will remove the sentence and reformat the following sentence.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC89 -
AC90: 'Reply on RC4.22', Leigh-Ann Smit, 02 Feb 2026
Line 190- the definition of bioturbation should be at the start of the section
We will move the definition of bioturbation to the beginning of the paragraph.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC90 -
AC91: 'Reply on RC4.23', Leigh-Ann Smit, 02 Feb 2026
Line 195- Later in the text, burrow architecture becomes key context in differentiating impacts. Different architectures should be defined here
We will add a sentence explaining the different types of burrow architecture (e.g simple burrow, branched burrow, chambered burrow, multi entrance burrow).
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC91 -
AC92: 'Reply on RC4.24', Leigh-Ann Smit, 02 Feb 2026
Line 201- What are destabilising and stabilising organisms? Some description should be given to these categories
Destabilizing organisms are those that disturb and mix sediment, making it looser and more erodible. Typically burrowing organisms. These include molluscs and crustaceans for example. Stabilising organisms are those that bind and compact sediment, increasing its resistance to erosion. These include organisms such as tube building polychaetes for example who produce mucus to bind the sediment. We will add a description to the text to give more context.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC92 -
AC93: 'Reply on RC4.25', Leigh-Ann Smit, 02 Feb 2026
Line 210- The outline of this sentence can help organize the following paragraphs. Physical, chemical, and biological impacts should each be described separately throughout each of the following sections
We will ensure these 3 aspects are covered in the sections that follow (accretion, erosion, sediment transport and deposition), and add information if it is not covered already.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC93 -
AC94: 'Reply on RC4.26', Leigh-Ann Smit, 03 Feb 2026
Line 212- This paragraph starts to get specific, and most of these points could be worked into the following sections, where this level of detail is more warranted
We will remove the paragraph and incorporate the relevant sentences into the following sections, where this level of detail is more appropriate.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC94 -
AC95: 'Reply on RC4.27', Leigh-Ann Smit, 03 Feb 2026
Line 241- Figure 3 has strong potential, and would benefit from stronger text support and some figure reorganization. Stronger text support: currently the text is organised using impacts of bioturbation, while figure 3 is organised by types of bioturbation and mechanisms that drive the impacts. This makes the figure difficult to follow along with through the text. Organising the figure in the same structure as the text would improve legibility. Figure reorganisation: the lower diagram doesn’t line up visual representations in the upper diagram, and this makes the figure difficult to follow. There are also some elements that are small/difficult to read in print. Also, the box for bioturbation impact #5 is blue, while the rest are grey- I don’t see an explanation for this in the caption
We will focus on incorporating Figure 3 more coherently in the text. Based on comment 2 from Reviewer 1, we will be adding a table which will include physical mechanisms of burrow construction and maintenance, looking at compaction, bioirrigation and sediment displacement. This will also include how these processes differ across different taxa and substrates. This will improve the link between the figure and the text. We will also reorganize the figure so that it follows the same structure as the text to ensure legibility. We will resize the elements in the figure so that they are legible. The blue box associated with #5 represents carbon and the grey shading represents sediment. We will include this explanation to the figure caption.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC95 -
AC96: 'Reply on RC4.28', Leigh-Ann Smit, 03 Feb 2026
Line 243- This is the opening paragraph of section on bioturbation, yet doesn’t mention bioturbation. This information should be moved, perhaps to a ‘tidal wetland geomorphology’ section in the introduction
We agree, we will relocate the first paragraph, lines 243-254, to the introduction section.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC96 -
AC97: 'Reply on RC4.29', Leigh-Ann Smit, 03 Feb 2026
Line 244- This is more of a focus for submerged vegetation such as seagrasses, which are not considered in this review. Regular inundation dramatically reduces salt marsh photosynthetic capacity
Thank you, we will remove that part of the sentence since it doesn’t make sense in the context of this review.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC97 -
AC98: 'Reply on RC4.30', Leigh-Ann Smit, 03 Feb 2026
Line 256- “extended periods and across surrounding areas” is vague
By “extended periods” we refer to long term effects, occurring over seasonal or multi-year timescale, not just over a single tidal cycle. By “surrounding areas” we refer to km-scale impacts, not just the immediate vicinity of the organism’s burrow. We will clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC98 -
AC99: 'Reply on RC4.31', Leigh-Ann Smit, 03 Feb 2026
Line 263- Are these bioturbators?
No, microphytobenthos (MPB) are not bioturbators. The inclusion of MPB in the text was meant as a comparison between some crab species that produce extracellular polymetric substances, such as mucus, which stabilises the sediment, in the same way that MPB stabilises the sediment. Later in the manuscript it is also mentioned that some bioturbators, such as crabs, consume MPB which then contributes to the destabilisation of sediment.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC99 -
AC100: 'Reply on RC4.32', Leigh-Ann Smit, 03 Feb 2026
Line 268- This doesn’t quite fit in the accretion section. Perhaps this is a better fit in the carbon sequestration, or in a soil chemistry section
The sentence “Low to moderate levels of bioturbation can be beneficial to primary productivity (Kristensen et al. 2008)” will be moved to the introduction section under the subheading Key carbon processes.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC100 -
AC101: 'Reply on RC4.33', Leigh-Ann Smit, 03 Feb 2026
Line 281- Slightly repetitive in this placement-this could be moved earlier
The line “the composition of crabs has the potential to influence ecosystems differently (Agusto et al. 2021)” will be moved to the paragraph above where it is currently positioned.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC101 -
AC102: 'Reply on RC4.34', Leigh-Ann Smit, 03 Feb 2026
Line 285- This statement doesn’t match the table and seems to be specific to this study-if so, that should be clarified
Yes, the statement “In salt marshes, however, bioturbation had no significant effect on changes in surface elevation” is specific to the study. We will ensure we clarify this in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC102 -
AC103: 'Reply on RC4.35', Leigh-Ann Smit, 03 Feb 2026
Line 322- is this always true? Earlier it was written that burrowers can enhance sediment cohesion
The statement “Burrowing activities weaken mud and clay banks in tidal marshes” is generally true. There are exceptions where some species e.g. Sesarma reticulatum secrete mucus which they coat their burrow walls with, which in turn can locally increase sediment cohesion. We will make this clearer in the text.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC103 -
AC104: 'Reply on RC4.36', Leigh-Ann Smit, 03 Feb 2026
Line 361- this is somewhat repetitive with sediment accretion and erosion above
We will review section 1.4.3 (sediment transport and deposition) to remove redundant content and make sure the information presented in the section provides additional insight.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC104 -
AC105: 'Reply on RC4.37', Leigh-Ann Smit, 03 Feb 2026
Line 415- Paragraph should have a more descriptive topic sentence
We will ensure the first sentence is more descriptive. We will alter it along the lines of “Consumers play a critical role in regulating carbon cycling, by directly and indirectly altering their physical environment, including primary production and sediment processes”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC105 -
AC106: 'Reply on RC4.38', Leigh-Ann Smit, 03 Feb 2026
Line 441- Could this also aerate and increase decomposition of previously stable C?
Yes, it could, we will include this information to that sentence, referencing Wang et al. 2010, who showed that bioturbation by crabs can promote sediment and carbon turnover (Wang, J. Q., Zhang, X. D., Jiang, L. F., Bertness, M. D., Fang, C. M., Chen, J. K., Hara, T., and Li, B.: Bioturbation of Burrowing Crabs Promotes Sediment Turnover and Carbon and Nitrogen Movements in an Estuarine Salt Marsh, Ecosystems, 13, 586– 599, https://doi.org/10.1007/s10021-010-9342-5, 2010.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC106 -
AC107: 'Reply on RC4.39', Leigh-Ann Smit, 03 Feb 2026
Line 488- This can also be from biological drivers
We agree that surface elevation can also increase as a result of biological drivers. We will edit the text to make it clear that both physical and biological processes contribute.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC107 -
AC108: 'Reply on RC4.40', Leigh-Ann Smit, 03 Feb 2026
Line 490- Repetitive with above
We will remove lines 490-495 as they are redundant.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC108 -
AC109: 'Reply on RC4.41', Leigh-Ann Smit, 03 Feb 2026
Line 502- Winter freeze occurrence
We will include winter freeze occurrence to the sentence, it will read “Mangroves are specifically vulnerable to changes in temperature and precipitation regimes, because the distribution range globally is linked to sea surface temperature and the occurrence of winter freezes”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC109 -
AC110: 'Reply on RC4.42', Leigh-Ann Smit, 03 Feb 2026
Line 504- The reference is missing
Thank you, the reference is: Tomlinson, P.B., 1999. The Botany of Mangroves. Cambridge University Press, Cambridge, United Kingdom. We will be sure to add it to the reference list.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC110 -
AC111: 'Reply on RC4.43', Leigh-Ann Smit, 03 Feb 2026
Line 539- “Increases in drainage density is necessary to manage the expanding tidal prism and effectively drain the marsh surface to prevent waterlogging”- Abrupt shift in topic
We will improve the link between the sentence “Increases in drainage density is necessary to manage the expanding tidal prism and effectively drain the marsh surface to prevent waterlogging” and crabs. The sentence will change to “Crab activity at tidal creeks can effectively increase drainage density, helping to manage the expanding tidal prism and prevent waterlogging on the marsh surface (Farron et al. 2020)”.
Citation: https://doi.org/10.5194/egusphere-2025-4749-AC111
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This manuscript presents an overview of the effect of bioturbation on tidal wetlands. While the topic is of great interest, the material is poorly organized, making it difficult for readers to develop a general understanding of the subject. As it stands, the manuscript reads more like a collection of disparate results from the literature rather than a cohesive synthesis. Below, I outline major points for improvement, along with specific suggested edits.
Major Points for Improvement:
Some additional minor edits are noted in the text.