the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
| 27 May 2025
The most complete Holocene peat record from Central Europe: multi-proxy reconstruction of postglacial wetness changes and climate events from Linje peatland, Poland
Abstract. This study investigates the vegetation and hydrological dynamics of Linje peatland in northern Poland during the past 11,500 years by integrating testate amoeba and plant macrofossil analyses. The Linje profile is currently the only complete Holocene peat record in Central Europe and offers valuable insights into long-term climate variability and its ecological consequences for peatland ecosystems. The results reveal significant changes in peatland wetness and vegetation driven by 20 autogenic processes, climatic fluctuations and anthropogenic influences. Major bryophyte species turnovers occurred around 11,200, 10,350, 8,200, 7,500, 5,500, 600, and 450 cal. BP often coinciding with declines in Archerella flavum and Hyalosphenia papilio abundances. Both proxies indicate a marked period of drier conditions between 7,600 and 6,800 cal. BP corresponding with the Holocene Thermal Maximum. Additionally, testate amoeba data suggest further disturbances at approximately 3,050, 2,000, and 200 cal. BP, and the latter, linked to a permanent shift in species composition, implies lasting changes to peatland 25 conditions caused by intensified human activity. Around the end of the Early Holocene, most species turnovers and disturbances began to align with Holocene Rapid Climate Change (RCC) events.
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Notice on discussion status
The requested preprint has a corresponding peer-reviewed final revised paper. You are encouraged to refer to the final revised version.
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Preprint
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The requested preprint has a corresponding peer-reviewed final revised paper. You are encouraged to refer to the final revised version.
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Interactive discussion
Status: closed
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RC1: 'Comment on egusphere-2025-2087', Anonymous Referee #1, 18 Jun 2025
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AC2: 'Reply on RC1', Eliise Poolma, 22 Aug 2025
We would like to sincerely thank the reviewer for the very helpful comments and suggestions that have significantly improved the quality and clarity of our manuscript. We have carefully considered all points raised and revised the text accordingly. In the specific replies below, we address each of the comments. We have expanded several explanations, improved clarity and language, and updated the reference list with more recent publications (including the references suggested by the reviewer). Regarding the NMDS ordination, we acknowledge the reviewer’s concerns, and we intend to revise this analysis accordingly (details provided in our specific replies). All technical corrections suggested by the reviewer have been implemented directly in the manuscript.
L35–38: Clarify what is meant by "main peat-forming species"; consider specifying taxa but also highlight that Sphagnum is not a prerequisite for peat formation”
Thank you for this comment. This section is clarified as follows: “Among the various vascular plants and bryophytes that contribute to peat formation, Sphagnum mosses stand out as particularly valuable in situ biological proxies in paleoecological research (Lamentowicz et al., 2019b; Ronkainen et al., 2014; Rydin et al., 2006; Clymo, 1984).”
L38: Include the citation: “https://doi.org/10.1016/0169-5347(95)90007-1” to support the ecological role of Sphagnum here. It might be helpful to add a few lines summarising its key traits.
Reference included and key traits are summarised in the following way: “The unique morphological, physiological, and biochemical traits of Sphagnum mosses make them effective ecosystem engineers, capable of building peat from their dead tissue while creating acidic, nutrient-poor, and waterlogged conditions. In doing so, they suppress the growth of vascular plants, further reinforcing conditions that favour their own dominance and continued growth (van Breemen, 1995; Turetsky et al., 2025).”
L34–36: Add a few lines discussing the value of Sphagnum in palaeoecology—it has many unique properties that make it valuable (e.g., decay resistance, vertical growth), but it's not essential for peat to form.
Regarding the two previous comments. We included the citation, emphasised that Sphagnum is not the only peat former, and added a section of key traits of Sphagnum as follows: “Among the various vascular plants and bryophytes that contribute to peat formation, Sphagnum mosses stand out as particularly valuable in situ biological proxies in paleoecological research (Lamentowicz et al., 2019b; Ronkainen et al., 2014; Rydin et al., 2006; Clymo, 1984). This is due to their distinct ecological preferences, physiological limitations, slow vertical growth, and resistance to decay. Different Sphagnum species are confined to specific microhabitats that vary in pH, nutrient availability, and water table depth (Rydin et al., 2006). Species turnover is strongly linked to changes in peatland hydrology, as Sphagnum species differ in their sensitivity to moisture availability in the capitulum and cannot actively regulate water uptake or prevent water loss (Gong et al., 2020). Together, these traits make Sphagnum mosses reliable indicators of past environmental conditions, particularly fluctuations in water table levels. In the Northern Hemisphere, Sphagnum mosses are major contributors to peat accumulation, especially in ombrotrophic bogs and poor fens where they typically dominate (Halsey et al., 2000). In contrast, peat formation in rich fens is primarily driven by brown mosses (mainly the Amblystegiaceae family), and by the roots and rhizomes of vascular plants (Jaszczuk et al., 2024). The unique morphological, physiological, and biochemical traits of Sphagnum mosses make them effective ecosystem engineers, capable of building peat from their dead tissue while creating acidic, nutrient-poor, and waterlogged conditions. In doing so, they suppress the growth of vascular plants, further reinforcing conditions that favour their own dominance and continued growth (van Breemen, 1995; Turetsky et al., 2025). Sphagnum communities growing on sufficiently deep peat deposits are highly resistant to water table changes, retaining their capacity to store carbon and continue growth even during drought conditions (Lamentowicz et al., 2019b; Marcisz et al., 2020a; Moore et al., 2021).”
L42–44: The claim that testate amoebae respond more quickly to water table changes than plants is questionable. There's an issue of circularity: vegetation and amoebae (from the same core) are not truly independent- vegetation can affect testate amoeba communities. Also, at the timescales typical of palaeoecological analysis, the resolution is often too coarse to capture these response-time differences (e.g., a 1 cm slice may represent decades or even a century or more).
It is true that testate amoebae have a dependency on vegetation. However, testate amoebae need much less time to change the community than plants. For example, van der Knaap et al. (2011) by sampling the peat in 2 mm slices that represented annual resolution, show that, while Sphagnum fuscum dominated, many different shifts in the testate amoebae community occurred .
van der Knaap, W.O., Lamentowicz, M., van Leeuwen, J.F.N., Hangartner, S., Leuenberger, M., Mauquoy, D., Goslar, T., Mitchell, E.A.D., Lamentowicz, Ł., Kamenik, C., 2011. A multi-proxy, high-resolution record of peatland development and its drivers during the last millennium from the subalpine Swiss Alps. Quaternary Science Reviews 30, 3467–3480. https://doi.org/10.1016/j.quascirev.2011.06.017
We changed the sentence in the manuscript in the following way: “While testate amoebae and plants are ecologically interconnected, testate amoebae communities can adjust species composition over shorter timescales due to their short life cycles and high turnover rates, whereas mosses and many vascular plants tend to respond more gradually to hydrological changes (Marcisz et al., 2014, Väliranta et al., 2012)”
L56: Be more specific about how anthropogenic land use change has damaged Holocene peat records, e.g., drainage, cutting, cultivation.
This section has been specified: “Full Holocene peat cores from Europe are rare because of the temporal variation in peatland succession (Ruppel et al., 2013; Stivrins, 2025) as well as drainage-based economic use ( Finlayson and Spiers, 1999; United Nations Environment Programme, 2022; Verhoeven, 2014).”
L69–72: It would help to briefly explain why this site is important for studying climate response. Who will this be useful for? You could add something like: “to better understand past ecosystem responses to climate change.”
Thank you for pointing this out. Specification added as suggested.
L94: There's missing information here: where were the radiocarbon dates processed? What material was dated? What pre-treatment methods were used? Are these your own dates or from an earlier study? If the former they should be in the results, if the latter a citation is needed.
All radiocarbon dates presented in this study are original and were obtained for this research. The dated material and laboratory codes are listed in Table 1, which we kept in the Materials and Methods, as the age-depth model based on these dates follows in the Results. All samples were processed at the Poznań Radiocarbon Laboratory (Poland) following their standard pre-treatment procedures for plant macrofossil material. As this is a well-established and routinely applied protocol in radiocarbon dating, we did not provide further methodological detail in the text.
L107: Why did you use both 5 and 10 cm³ samples? Which depths used which volume?
We used samples of varying volume because the macrofossil study had two goals: the primary one was analysing plant macrofossils, originally planned at a 10 cm resolution, and the secondary goal was to recover as many subfossil Betula nana leaves as possible for a separate study. To capture finer-scale changes of interest, especially in key sections of the profile, we increased the resolution to every 5 cm. The step of the samples (either 5 cm or 10 cm) is visible in the supplementary dataset. The studied volume will also be added to the table.
L110–135: More methodological detail needed for the testate amoeba work: what magnifications were used? Which identification guides? What was the minimum test count? And just to clarify, was a stereomicroscope also used for this part? I would usually use a high-powered microscope only.
All these information have already been included in the original manuscript – see preprint pdf, page 5 lines 125-134. Only a light microscope has been used for the analysis.
L148: It would be useful to include some detail on the transfer function model used: what type of model, what is the R² and RMSE, and maybe clarify that it was run using Rioja (and cite it).
The information about the transfer function model has already been included in the original manuscript (see the section explaining statistical methods). The R2 and RMSE of the model are explained in detail in the cited publication - Amesbury et al. 2016.
L160: The structure is a bit confusing-it feels like you have two interpretation sections: one here and one later. I’d consider merging these two sections into a single “results and interpretation” section, where each environmental change is described and interpreted in turn. This would make your paper much clearer.
We thank the reviewer for this suggestion. Our intention was to first present proxy-related changes in detail with ecological interpretation and then place the changes into a broader climate-related discussion with comparison to other studies in the next chapter. Thus we would like to keep the structure, but we have also carefully reviewed and revised the Results section so that no climatological interpretation would appear there.
L164: I’d rephrase “should reflect” as something like “we lack the chronological control to detect short-term differences in accumulation rates.” Also, instead of “treated with care,” maybe say “interpreted with caution due to this limitation.” The interpretation you give later seems reasonable, so no major concern there.
Rephrased according to the recommendation: “For the topmost 100 cm we lack the chronological control to detect short-term differences in accumulation rates. This section should thus be interpreted with caution due to this limitation, as it is based on one date and it does not reflect an exponential depth-age relationship as it should have with well-developed acrotelm and catotelm layers (Marcisz et al., 2015).”
L175: Rather than just referring the reader to the figures, I’d expand a little here. Which species are dominant? Any obvious community shifts or interpretive points worth flagging?
Thank you for this suggestion. The major changes in the proxies are outlined in the beginning of the section 5.4 (“Interpreting the proxy indicated changes”). However, we agree that moving this information where the reviewer suggests might increase clarity and flow. This will be revised in the manuscript.
L189: “Indicate the presence of a freshwater habitat” – in a peat-forming system that’s a given. Do you mean more like inundated conditions or open water? Could it have been a shallow lake phase?
Thank you for pointing this out - we did mean inundated conditions or a shallow water habitat. We also clarified this in the manuscript.
L198–199: Was the core sampled all the way to the base of the peat? Some interpretation of the factors that led to peat initiation might be interesting here if you can offer some explanation.
The core was sampled until the base of the peat, but we decided to handle the earliest stages this in a separate publication. These samples have not been analysed yet for plant macrofossils, thus we limit the discussion in this paper about the peat initiation.
L204: Replace “in the region” with “in the wider landscape” - “region” can imply a much broader area.
Replacement is made here and elsewhere.
L237: Define what is meant by “intermediate microform.”
By intermediate microform, we mean either a high lawn or low hummock microform, intermediate in the wet-dry gradient. S. medium has been found growing in hollows, lawns, and it can also extend to the hummock microform.
The section has been altered for clarity “Previously dominating wet-habitat Sphagnum subgen. Cuspidata is replaced by S. divinum/medium, which might suggest development of a drier microform (Blackford, 2000; Hassel et al., 2018).”
L261: G. discoides can indicate disturbance, yes—but it’s also found in quite stable systems. Maybe rephrase to something like: “can indicate fluctuating water tables, sometimes linked to disturbance.”
Thank you! The section is rephrased according to the suggestion
L287–288: What’s the significance of the D. rotundifolia seeds here? If this is their first appearance, could it reflect increasing nutrient deficiency?
The first occurrence of Drosera seeds is recorded in the sample dated to approximately 10,900 cal. BP. This likely reflects an increase in nutrient deficiency, as both D. rotundifolia and D. anglica, despite their slightly differing habitat preferences, are indicative of oligotrophic conditions. However, in the section spanning 4,000 to 600 cal. BP, I would not add any specific ecological interpretation to their presence, and thus this sentence is removed.
To the first appearance, we added to following section: “Differentiating between the seeds of the two Drosera species is difficult. D. rotundifolia tends to indicate lower pH and exhibits a broad ecological tolerance, whereas D. anglica is typically associated with wetter conditions and slightly higher pH levels (Banaś et al., 2023). Nonetheless, the presence of either insectivorous species implies increasing nutrient limitation in the peatland environment (Mauquoy and van Geel, 2007).”
L295: Have you got any evidence or references supporting this as an indicator of hydrological variability? What might be driving the pattern?
This interpretation was based on contrasting observation that included the co-occurrence of species indicative of both wetter and drier conditions, along with a decline in H. papilio and Sphagnum abundances. However, due to the lack of supporting references, this section has been revised as follows:
„Around 2,000 cal. BP, drier conditions are inferred from an increase in B. indica and A. seminulum abundances, accompanied by a decline in H. papilio. This shift corresponds to a reconstructed water table drop to approximately 12 cm. The timing also coincides with low Sphagnum percentages and an increase in decomposed organic matter, collectively indicating increased hydrological instability.”
L292: There seems to be a mismatch between dry-indicator testate species and wetter Sphagnum types - could this be a taphonomic/preservation issue or something else?
Around 2,000 cal. BP, the presence of moderately wet Sphagnum species contrasts with testate amoeba indicators, suggesting drier conditions. However, the overall Sphagnum percentage is low, accompanied by increased amounts of decomposed organic matter and a rise in Ericaceous remains. These patterns likely reflect a relatively rapid drying event to which the Sphagnum community was unable to respond immediately. As Sphagnum cover later recovers, a compositional shift toward slightly drier conditions is evident, marked by the replacement of Sphagnum sect. Cuspidata with S. divinum/medium.
L319: Do you think the uppermost WTD reconstructions are affected by preservation bias (e.g., due to preservation/dominance of idiosomic species as you state)?
No, we do not think that the preservation bias is an issue here. We observed often in the previous studies that presence of idiosomic species in the cores is related to landscape opening and deforestations by local communities (see e.g., Lamentowicz et al. 2013 Journal of Quaternary Science; Gałka et al. 2014 Quaternary International; Marcisz et al. 2025 Scientific Reports; Lamentowicz et al. 2020 Scientific Reports; Lamentowicz et al. 2019 Quaternary Science Reviews). Moreover, the hypothesis claiming the decomposition of idiosomic shells in peat profiles is based on a single experimental study from almost two decades ago (Swindles and Roe 2007 Paleo3) and it has not been confirmed by any other study. Therefore, we think it is speculative.
L323: I’m not convinced it makes sense to compare the testate amoeba data with the DWT, since the DWT is derived from the testate communities. A better comparison might be to the plant macrofossils. Also, the NMDS is underused, it’s not really integrated into the discussion. You could either expand the interpretation or consider removing it altogether.
Thank you for addressing this important issue. We agree that the ordination is currently underrepresented in the text. We also consider moving it to the supplementary materials. To avoid the problem of circularity in comparing testate amoebae with DWT, we will revise the NMDS by removing the DWT variable and focusing instead on the relationships between the biological proxies.
L344–350: Not clear what this NMDS ordination really adds. Fire is barely discussed in the text, yet macrocharcoal plays a role in this plot. Consider dropping or restructuring this part unless you can tie it in more meaningfully. The discussion of species clustering is also underdeveloped, and could merit some statistical analysis and inclusion in the discussion.
Thank you! We will also remove the macrocharcoal variable from the NMDS ordination and expand the discussion about the species clustering together on the plot.
L352–355: The note on hummock species surviving fire is interesting, maybe bring that up earlier in the discussion? Fire is not really a central theme of this paper but there are some macrocharcoal peaks.
Thank you for the suggestion. As mentioned before by the reviewer, fire plays a minor role in this paper and our dataset, thus we have decided to omit this part from the discussion.
Fig. 7: Would it be helpful to expand on how the wetness changes relate to known RCC phases? You could also consider adding the lake Żabieniec record directly into the figure for comparison?
Thank you for this suggestion. While we agree that linking wetness changes to known Rapid Climate Change (RCC) phases could be informative, we note that RCC intervals are generally long and characterized by fluctuating climate conditions rather than clearly defined wet or dry events. In our record, this is expressed more through phases of species turnover and hydrological instability/disturbances rather than straightforward wetness shifts. For this reason, we consider it more appropriate to expand on these connections in the main text rather than directly in the figure (where the reconstructed wetness changes from our proxies are already presented). We will also make this point more explicit in the text. As recommended, we will add the Lake Żabieniec record to Figure 7.
Fig. 7 caption: Clarify—by “disturbances” do you mean anthropogenic or climatic?
Mostly climatic but since a clear anthropogenic disturbance is visible, we will include both forms of disturbance.
L397–400: This might tie in with the chironomid-based reconstructions, could be worth pointing that out. Are we seeing cooling or drying reflected here?
Thank you for pointing this out. We do mention the cooling indicated by the chironomid record later in this paragraph (lines 404-405), but we agree that this section could benefit from some restructuring to improve clarity. The Linje record suggests either a dry event or at least a period with water table fluctuations. However, based solely on our record, it is not possible to determine whether the decomposition and bryological shift were driven by cooling or warming. Other regional records, such as varves (Zander et al., 2024) and chironomids (Kotrys et al., 2020) show contrasting results during this period.
Pg. 17, Discussion: I think you could say more about the implications of this work for future climate change. What can we learn from these past dynamics?
Thank you for this valuable suggestion. We agree that our study can be better placed in the context of future climate change. Our record shows that for millennia, the peatland was resilient to natural climate variability, maintaining peat accumulation despite repeated hydrological and vegetational shifts. However, following drainage in recent centuries, the system has not shown recovery, which highlights the limits of resilience once strong anthropogenic pressures are introduced. We will expand this in the “Conclusions”.
L401: It looks like there’s a phase of terrestrialisation here, possibly linked to the 8.2 ka event? It’s worth noting that this event was quite drawn-out in some records, with a lead-up to the main climate shift.
Thank you for this comment. We interpret the changes around this section as a transition from rich fen to poor fen and we agree that after the disturbance between 8500-8300 there is a shift to drier conditions. We also acknowledge the possibility that these changes could be linked to broader hydrological variability around the 8.2 ka event, but given the resolution and nature of our data, we prefer to remain cautious in attributing the observed changes directly to the 8.2 ka event.
L404–405: What’s your take on the discrepancy here? Worth expanding.
As mentioned before - varve record (Zander et al., 2024) and chironomids (Kotrys et al., 2020) show contrasting results during this period. Our samples show high decomposition which is followed by a Sphagnum community shift to drier microform species (while the testate amoeba record does not show significant water table changes). Since dryness and decomposition could result for example from cold, more continental winters or dry and hot summers, then this remains to be clarified in future studies. There could be some seasonal variation that our resolution does not capture.
L428: The change in the TA record seems to be driven by increasing A. seminulum and A. militaris—you could discuss their ecology, especially A. militaris as a dry indicator.
A. militaris and A. seminulum are both typically present in Sphagnum-dominated peatlands – from bogs to poor-fens, and they tend to prefer drier conditions, water tables lower than 10 cm. Alabasta militaris is often present when water tables depth is somewhere between 10 to 20 cm. All these species are indicative of drier phases on the peatland and are commonly present/well preserved in peat cores.
L439: What might have caused the disturbance observed here?
The disturbance and species turnover in Sphagnum is likely caused by Mid-Holocene cooling like on many other peatlands in Europe. We clarify this in the manuscript.
L489: E. vaginatum is a peat-former, so its presence doesn’t necessarily indicate an interruption in peat accumulation, maybe rephrase.
Thank you for the comment. We rephrased as follows: “These sites display high water table fluctuations, growth of Eriophorum vaginatum and the disappearance of S. fuscum.”
L506: Could you explain how the process you're referring to works, for readers less familiar with it?
As mentioned before, we would like to pay less attention to fire activity in the revised manuscript version as it was not of importance for the development of Linje peatland, thus we will remove this statement from the revised manuscript and only point out that the hydrological shifts might be caused by increased human activity in the region.
L517: I’d be cautious about suggesting testate amoebae are simply better indicators than plants. They’re strongly influenced by vegetation themselves. Maybe include a bit of reflection on how this affects your interpretation.
It was not our intention to suggest that testate amoebae are better indicators than plants, as we compared only the differences in reaction speed. However, to avoid a misunderstanding, we removed the problematic phrase from the revised manuscript.
L520–521: Could be a good place to link back to the Swindles et al. (2019) Nature Geoscience paper on the drying of European peatlands. Your data seem to support it.
Thank you for the suggestion, we will include this reference and link it to the text.
L256: Linje is a well-studied site—do you know what the current average WTD is? Might be worth including here to support the reconstruction or provide some sense of its limitations.
The monitoring of Linje mire is ongoing. The recent analysis published by Słowinska et al. (2022 J of Biometeorology) show that water table changes are variable in different parts of the site. Especially the north-eastern part of the site is particularly vulnerable to drought as it lays on inland dune that effectively infiltrates the surface waters. Knowing the seasonal changes of hydrology of the site we cored in the most hydrologically stable central part of Linje to obtain the most objective information.
Section 5.4: This section is quite descriptive and doesn’t fully interpret the environmental changes discussed earlier. It would be better to merge it with the previous section into a single, interpretive narrative.
Thank you for the suggestion. We agree that some parts of the Section 5.4 could be merged with other parts of the manuscript. However, we wanted to separately emphasize that the observed hydrological shifts should be interpreted with some caution as
- a) the site experiences a unique microclimate now and this could have been a case in the past
- b) observed hydrological shifts are today and could have been in the past influenced by groundwater changes.
L550–551: This sounds like an interesting record, maybe include it more centrally in your interpretation or even in Fig. 7?
We thank for this suggestion but we would like to stay with the citation and discussion because of the different data structure in the cited publication.
L557–558: If this is a different core but shows synchronicity, that could support the representativeness of your record, at least for the past 2000 years.
Thank you for pointing this out. Yes, it is a different core and it is in a rather good agreement. We will emphasize this more in the revised manuscript.
L570: Could climate and hydrology be linked here? Or perhaps trophic shifts or mineral input played a role too- consider other possible drivers.
Thank you for pointing this out. We will expand this sentence while considering other possible drivers as well.
Citation: https://doi.org/10.5194/egusphere-2025-2087-AC2
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AC2: 'Reply on RC1', Eliise Poolma, 22 Aug 2025
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RC2: 'Comment on egusphere-2025-2087', Anonymous Referee #2, 20 Jun 2025
Thank you for the opportunity to review the article “The most complete Holocene peat record from Central Europe: multiproxy reconstruction of postglacial wetness changes and climate events from Linje peatland, Poland” by Poolma et al.,
The paper deals with the biological proxies, i.e., bryophytes, testate amoeba, microcharcoal and radiocarbon dating of the longest peat recorded in Linje mire, Poland. The topic itself is of interest and suitable for the journal. But the paper requires some confirmations, improvements, and modifications before publication. As indicated in the attached annotated pdf file more supporting evidence is required for some of the methodologies and discussions.
General comments:
The background of this study needs to be improved, and please clarify some issues. Several studies have been conducted in the Polish peatland in general, particularly in Linje mire, as stated in the background. Therefore, I can’t see clear aims or significance compared to the previous study. If the authors attempt to test the biological proxies using bryophytes and testate amoeba, therefore emphasizing and comparing the strengths and weaknesses with other proxies (e.g., pollen, geochemical, etc) should be explained. Holocene climatic fluctuations should be elaborated more, why these events give a significant contribution to further climate predictions, robustness interpretation, or new approaches to methodology. The interconnection between Holocene climatic fluctuations (stated in the background) is not clear with the Rapid Climate Change (in discussion). Other issues are references used are relatively “old”, but somewhat the authors state that paleoclimate reconstructions using peat deposits continue to develop recently. I suggest adding new published data and references to fill and extend the research gap. I would greatly appreciate it if the authors could provide chemical data to support the interpretations related to the change in peat chemistry and nutrient availability.
The methodology section should be enhanced and improved. Samples were taken from 2019; how the samples were obtained and treated needs to be clarified. A physical description of the core is needed to figure out the samples, since no representative photos are shown. Pre-treatment of samples should be clearly stated, especially in radiocarbon dating analysis. It is well known that there are some pre-treatments using Acid-Acid or Acid-Base-Acid methods to remove some inorganic materials. Other fundamental issues that should be taken into consideration are how they distinguish the macrofossils using plant parts into specific species of plants and how they count the particles. Representative photos of macrofossils, bryophytes, and charcoal should be given to help the readers figure out the data observed and obtained. It is not clear how the authors identify specific species of bryophytes using “fragments” solely, but somehow it is interesting too. The NMDS ordination approach for statistical analysis should be reconsidered (please find detailed comments). The NMDS ordination does not support the findings, or perhaps the authors should consider the plant macrofossils parameter to be included. In addition, a lot of “peatland chemistry” records were stated without any evidence of chemical analysis!
The discussion section is interesting, but the inconsistency using specific species or parameters should be considered. For example, drier conditions in Zone-3 were characterized by the presence of Salix herbacea and Empetrum nigrum and ericacous remains; however, drier conditions in Zone-5 and 6 were indicated by Sphagnum divinum and Sphagnum sect. Acutifolia species, primarily S. rubellum and S. fuscum. I suggest adding information on dry-related species and wet-related species in Figure 4. The terminology of “disturbance” and how they define this event is also unclear. Does it reflect the drier conditions, low nutrient availability, or other factors?? Moreover, the “disturbance” events should be further explored, either anthropogenic or climatic drivers. I am not sure that nutrient availability and change in chemistry of peats can be interpreted using change in biological proxy solely; these should be strengthened by chemical analysis such as C/N ratio, sulfur content, and other biogenic elements analysis. I suggest enhancing the interpretation using the correlation between DWT to the preservation/degradation particles to find out the controlling factor on peat preservation. Lastly, since there are a lot of paleoclimate records in Linje mire (reference therein), I would recommend adding a figure that shows the results and compares them to previous records, therefore we can see clearly the significance of this study. This integration could help the readers to have a broader point of view from local-regional scales.
I suggest to add some references related to this study that might help to enhanced better interpretations.
Evans et al., (2025) https://doi.org/10.1016/j.ejop.2025.126147
Sim et al., (2023; 2021) https://doi.org/10.1016/j.quascirev.2023.108020; https://doi.org/10.1016/j.ecolind.2021.108122
Swindles et al., (2020) https://doi.org/10.1016/j.ejop.2020.125693
Swindles et al., (2019) https://www.nature.com/articles/s41561-019-0462-z
Thank you
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AC3: 'Reply on RC2', Eliise Poolma, 22 Aug 2025
We would like to thank the reviewer for the careful reading of our manuscript and for providing a detailed set of comments. We acknowledge the concerns raised regarding the background, methodological detail, and clarity of some interpretations. In the revised manuscript and in our responses, we have outlined the significance of this study more clearly in comparison to earlier work from Linje peatland and other Polish peatlands. We have also added more recent references (including those suggested by the reviewer), and updated the literature to reflect current developments in peatland palaeoecology.
In addition, we have provided more detailed explanations of the plant macrofossil and testate amoeba methodologies, as well as clarifications on core retrieval, storage, subsampling, and radiocarbon dating. We have clarified the use of the term “disturbance” and agreed with the suggestions to revise the NMDS ordination approach.
While our interpretations are based on biological proxies, we fully agree that combining these with geochemical analyses would strengthen the study. Unfortunately, such data is not available at this stage but will be considered for future research.
Attached PDF provides detailed answers to all reviewer comments.
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AC3: 'Reply on RC2', Eliise Poolma, 22 Aug 2025
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RC3: 'Comment on egusphere-2025-2087', Anonymous Referee #3, 23 Jun 2025
Comments to “The most complete Holocene peat record from Central Europe: …“ by Poolma et al.
This discussion paper presents a study investigating a Holocene record from Poland. The science is sound and the results are very interesting so that this study should certainly be published. I am not an expert on Sphagnum and testate amoeba and the comments made above by reviewer 1 and 2 are quite detailed. Therefore I only add several details which I think could be considered.
As a non-expert, I would really like a figure showing examples for the testate amoeba and bryophytes encountered.
Line 42 etc.: While I agree generally, concerning vascular plants the differences inside this group are huge (1- and 2-year cycles vs. taxa with lifespans of decades).
Line 59: The record by Noryśkiewicz (2005) covers 106 pollen samples from a neighbouring site so that it may be interesting to mention it later too or even include it (important taxa/moisture indicators?) in one of the later figures (see below). It is only mentioned here and concerning Fig. 1.
Line 90 and following: Concerning the age model, I can completely understand that sample Poz.128079 was ignored, but I am not sure if Poz-128073 was automatically excluded by the used software or by the authors – maybe in case of Poz-128073, the exclusion could be explained in a little more detail.
Linbe 97 (if not changed): …as they were…
Line 92: I guess that ‘monolith’ means a profile, but I am not sure if it is referring to the core or an extra profile. It may be okay to use the term, but I at least was a little irritated.
Line 108: ‘fragment’ could be replaced by section or a similar term. ‘Fragment’ implies something is broken.
Line 111: ‘high-power microscope’? Please explain!
Line 143: For the non-expert, a table showing the assignments might help.
Figure 2: A little bit more of the approach should be given in the figure text. It does not name used software/interpolation methods.
Figures 3-5: In the preprint, the quality of the figures is not that good. In any case, genera/species names should be in italics in all figures (as they are in 3).
Line 180 and following: The results sometimes contain already interpretations – while some readers may prefer a clearer separation and to move interpretations into the discussion, I think this is okay this way. I suggest to avoid phrases like ‘we see’ or ‘we find’, but this is also a matter of taste.
Figure 6: While this figure is interesting and important, I wondered if it could be made more easily readable. Particularly for the outer taxa, more of the names could be given, and scale numbers are very small.
Line 356 and following: As implied above, I am a little confused that the results are (well) compared with over-regional records, including pollen records, but that the pollen record by Noryśkiewicz (2005) is not mentioned here. It may even be included in Fig. 7 (arboreal pollen vs. with revised age model?).
Line 385: According to Zurek (2005).
Line 525 and following: ‘water level’ may be confusing, I assume the depth to the water table is still meant here.
Line 534 and following: Some more details could be given which kind of anthropogenic influence could cause this.
Line 590-599: I think in several sentences, a ‘,’ should be placed before the ‘and’.
Citation: https://doi.org/10.5194/egusphere-2025-2087-RC3 -
AC1: 'Reply on RC3', Eliise Poolma, 21 Aug 2025
We would like to sincerely thank the reviewer for the time and effort dedicated to carefully evaluating our manuscript. Below, we provide a detailed response to all comments.
As a non-expert, I would really like a figure showing examples for the testate amoeba and bryophytes encountered.
Thank you for your interest! The species encountered in this core are very common and already well illustrated in existing resources, which is why we chose not to include photographs in the manuscript. We would, however, have provided figures if rare taxa or potentially new fossil species had been found. For high-quality images of testate amoebae, we recommend the Arcella website (http://www.arcella.nl). For bryophytes, detailed microscope images can be found in the British Bryological Society’s species finder (https://www.britishbryologicalsociety.org.uk/learning/species-finder/). The subfossil remains look very similar to the modern specimens illustrated there.
Line 42 etc.: While I agree generally, concerning vascular plants the differences inside this group are huge (1- and 2-year cycles vs. taxa with lifespans of decades).
We appreciate this observation and agree that vascular plants show considerable variation in life history strategies, ranging from annuals to perennials and clonal species. We have updated the text accordingly:
“While testate amoebae and plants are ecologically interconnected, testate amoebae communities can adjust species composition over shorter timescales due to their short life cycles and high turnover rates, whereas mosses and perennial vascular plants tend to respond more gradually to hydrological changes (Marcisz et al., 2014; Väliranta et al., 2012).”
Line 59: The record by Noryśkiewicz (2005) covers 106 pollen samples from a neighbouring site so that it may be interesting to mention it later too or even include it (important taxa/moisture indicators?) in one of the later figures (see below). It is only mentioned here and concerning Fig. 1.
Noryśkiewicz (2005) presents only a preliminary description of pollen zones without age estimates (reported only as depths). Unfortunately, we did not find a way to meaningfully integrate these results into our figures or analyses.
Line 90 and following: Concerning the age model, I can completely understand that sample Poz.128079 was ignored, but I am not sure if Poz-128073 was automatically excluded by the used software or by the authors – maybe in case of Poz-128073, the exclusion could be explained in a little more detail.
We did preliminary calculations of the model, including all dates, Both dates Poz-128079 and Poz-128073 revealed lack of compatibility to the model. By applying each radiocarbon date to the model calculation it was not possible to produce a valid chronology. Poz-128073 was not automatically excluded but after testing models:
Resolving order: Finding R_Date Poz-128079. After excluding Poz-128079, the agreement of age-depth model (Amodel) was still equal to 0. However, both dates Poz-128074 and Poz-128073 revealed the lack of compatibility with the model (Agreement of dates to the model equal to 1%). In next step, we calculated two models X) with dates Poz-128079 and Poz-128073 excluded, and Y) with dates Poz-128079 and Poz-128074 and we compared Amodel of X) and Y). Applying the same parameters i.e P_Sequence(k0=1, interpolation=2, log10(k/k0=1) the model X revealed Amodel=66%, whereas the model Y revealed Amodel=7%. Therefore, we selected model X) as one used as a basis for absolute chronology applied for this study (profile) as it is above recommended minimum equal to 60 % (Bronk Ramsey, 2008).
To the manuscript we added a short additional explanation: “Dates Poz-128079 and Poz-128073 were distinctly younger than the neighbouring dates. Preliminary model runs, both with the full set of dates included and with only one problematic date excluded, showed that these two ages had very poor agreement with the model. Therefore, both were excluded from the final age–depth model.”
Linbe 97 (if not changed): …as they were…
Change made in the revised manuscript version
Line 92: I guess that ‘monolith’ means a profile, but I am not sure if it is referring to the core or an extra profile. It may be okay to use the term, but I at least was a little irritated.
Thank you for pointing this out. To clarify, two parallel peat profiles were originally extracted at a distance of 0.5 m. However, in this study, we analysed only one of these profiles. To avoid confusion, we have revised the text accordingly.
Line 108: ‘fragment’ could be replaced by section or a similar term. ‘Fragment’ implies something is broken.
Change made in the revised manuscript version
Line 111: ‘high-power microscope’? Please explain!
Thank you for pointing this out. We mean light microscope.
Line 143: For the non-expert, a table showing the assignments might help.
Thank you for the suggestion, we have added a table to the revised manuscript that clarifies the assignment of bryophytes to the functional group.
Figure 2: A little bit more of the approach should be given in the figure text. It does not name used software/interpolation methods.
Thank you for pointing this out, we have added the missing information to the figure caption in the revised manuscript.
Figures 3-5: In the preprint, the quality of the figures is not that good. In any case, genera/species names should be in italics in all figures (as they are in 3).
The figures quality decreased after the submission; we guess it is the case of preprint repository. We have the figures in better quality and can provide them for the final manuscript version. We will make sure all taxa names are italicised on the figures, thank you for pointing this out.
Line 180 and following: The results sometimes contain already interpretations – while some readers may prefer a clearer separation and to move interpretations into the discussion, I think this is okay this way. I suggest to avoid phrases like ‘we see’ or ‘we find’, but this is also a matter of taste.
Thank you for this comment. We acknowledge that some ecological interpretation appears in the Results section, particularly where it is closely tied to the description of proxy data. We chose this structure because we find that presenting ecological interpretation alongside the proxy changes improves clarity for the reader. In contrast, the broader climatological interpretation and comparison with other studies are in the Discussion. We have carefully reviewed the text to ensure a clearer distinction between zone descriptions and interpretation, and we have also revised the phrases mentioned.
Figure 6: While this figure is interesting and important, I wondered if it could be made more easily readable. Particularly for the outer taxa, more of the names could be given, and scale numbers are very small.
We will improve the readability of the figure as suggested
Line 356 and following: As implied above, I am a little confused that the results are (well) compared with over-regional records, including pollen records, but that the pollen record by Noryśkiewicz (2005) is not mentioned here. It may even be included in Fig. 7 (arboreal pollen vs. with revised age model?).
Thank you for this suggestion. We agree that including the study by Noryśkiewicz (2005) would indeed strengthen the regional comparison. However, as this publication only provides a general preliminary description of the pollen zones without references to age estimations (only depths), we were unfortunately not able to incorporate it further into our manuscript or figures. We will acknowledge this limitation in the revised text.
Line 385: According to Zurek (2005).
Change made in the revised manuscript version
Line 525 and following: ‘water level’ may be confusing, I assume the depth to the water table is still meant here.
Thank you for pointing this out – the phrase “water level” was replaced by “water table” or “depth to the water table” in the manuscript depending on the context.
Line 534 and following: Some more details could be given which kind of anthropogenic influence could cause this.
Thank you for pointing out this inconsistency. The Sphagnum species turnover observed 200 years ago could have both climatic (Little Ice Age) as well as anthropogenic (deforestation etc.) reasons. This is specified in the revised manuscript.
Line 590-599: I think in several sentences, a ‘,’ should be placed before the ‘and’.
Change made in the revised manuscript version
Citation: https://doi.org/10.5194/egusphere-2025-2087-AC1
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AC1: 'Reply on RC3', Eliise Poolma, 21 Aug 2025
Interactive discussion
Status: closed
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RC1: 'Comment on egusphere-2025-2087', Anonymous Referee #1, 18 Jun 2025
Review of The most complete Holocene peat record from Central Europe: multi-proxy reconstruction of postglacial wetness changes and climate events from Linje peatland, Poland; Eliise Poolma et al. Climate of the past
- General comments
This manuscript presents a new, complete Holocene reconstruction of environmental change from Linje mire in Poland, currently the only complete peat record for this epoch in Central Europe. This alone makes this study a valuable and significant contribution, well aligned with the scope of Climate of the Past.
The title clearly reflects the contents of the manuscript, although it does not explicitly mention the anthropogenic changes observed towards the end of the record, this constitutes a relatively small (albeit significant) portion of the study and does not detract from the overall clarity or relevance of the title. The abstract provides a concise and accurate summary of the study
The study introduces new data, including a robust age-depth model based on 14 AMS radiocarbon dates, as well as stratigraphic records of botanical peat composition and testate amoebae. These are effectively used to infer hydrological variability for the site throughout the Holocene. The interpretations and conclusions are well supported by the presented data, and the study includes a thorough comparison with other palaeoecological records. However, in some instances, the manuscript could benefit from a more detailed explanation of observed changes, better contextualised within the broader peatland literature, and could benefit from some re-structuring. Doing so would further highlight the significance of this site and the value of the record. Methodologically, the study follows standard and appropriate protocols, which are clearly described and allow for reproducibility, although the NMDS ordination compares testate amoeba communities with water table reconstructions derived from the same data, which is inappropriate as obviously there will be a relationship between the two records (it is interesting that the relationship is quite weak). This analysis does not contribute much to the main discussion/findings and should be omitted or changed, perhaps to better relate plant macrofossils to the DWT reconstruction for interpretation (See specific comments). The manuscript is mostly well referenced throughout (see technical comments), with an appropriate number of citations, although several references are relatively old, and one is unpublished or still under review, this is not a major objection.
The original contributions of the authors are clearly stated. The language is generally fluent and technically accurate, though there are minor stylistic and grammatical issues in places. Suggestions have been made to improve clarity, accuracy and flow in certain sections (see technical corrections).
Specific comments
L35–38: Clarify what is meant by "main peat-forming species"; consider specifying taxa but also highlight that Sphagnum is not a prerequisite for peat formation”
L38: Include the citation: “https://doi.org/10.1016/0169-5347(95)90007-1” to support the ecological role of Sphagnum here. It might be helpful to add a few lines summarising its key traits.
L34–36: Add a few lines discussing the value of Sphagnum in palaeoecology—it has many unique properties that make it valuable (e.g., decay resistance, vertical growth), but it's not essential for peat to form.
L42–44: The claim that testate amoebae respond more quickly to water table changes than plants is questionable. There's an issue of circularity: vegetation and amoebae (from the same core) are not truly independent- vegetation can affect testate amoeba communities. Also, at the timescales typical of palaeoecological analysis, the resolution is often too coarse to capture these response-time differences (e.g., a 1 cm slice may represent decades or even a century or more).
L56: Be more specific about how anthropogenic land use change has damaged Holocene peat records, e.g., drainage, cutting, cultivation.
L69–72: It would help to briefly explain why this site is important for studying climate response. Who will this be useful for? You could add something like: “to better understand past ecosystem responses to climate change.”
L94: There's missing information here: where were the radiocarbon dates processed? What material was dated? What pre-treatment methods were used? Are these your own dates or from an earlier study? If the former they should be in the results, if the latter a citation is needed.
L107: Why did you use both 5 and 10 cm³ samples? Which depths used which volume?
L110–135: More methodological detail needed for the testate amoeba work: what magnifications were used? Which identification guides? What was the minimum test count? And just to clarify, was a stereomicroscope also used for this part? I would usually use a high-powered microscope only.
L148: It would be useful to include some detail on the transfer function model used: what type of model, what is the R² and RMSE, and maybe clarify that it was run using Rioja (and cite it).
L160: The structure is a bit confusing-it feels like you have two interpretation sections: one here and one later. I’d consider merging these two sections into a single “results and interpretation” section, where each environmental change is described and interpreted in turn. This would make your paper much clearer.
L164: I’d rephrase “should reflect” as something like “we lack the chronological control to detect short-term differences in accumulation rates.” Also, instead of “treated with care,” maybe say “interpreted with caution due to this limitation.” The interpretation you give later seems reasonable, so no major concern there.
L175: Rather than just referring the reader to the figures, I’d expand a little here. Which species are dominant? Any obvious community shifts or interpretive points worth flagging?
L189: “Indicate the presence of a freshwater habitat” – in a peat-forming system that’s a given. Do you mean more like inundated conditions or open water? Could it have been a shallow lake phase?
L198–199: Was the core sampled all the way to the base of the peat? Some interpretation of the factors that led to peat initiation might be interesting here if you can offer some explanation.
L204: Replace “in the region” with “in the wider landscape” - “region” can imply a much broader area.
L237: Define what is meant by “intermediate microform.”
L261: G. discoides can indicate disturbance, yes—but it’s also found in quite stable systems. Maybe rephrase to something like: “can indicate fluctuating water tables, sometimes linked to disturbance.”
L287–288: What’s the significance of the D. rotundifolia seeds here? If this is their first appearance, could it reflect increasing nutrient deficiency?
L295: Have you got any evidence or references supporting this as an indicator of hydrological variability? What might be driving the pattern?
L292: There seems to be a mismatch between dry-indicator testate species and wetter Sphagnum types - could this be a taphonomic/preservation issue or something else?
L319: Do you think the uppermost WTD reconstructions are affected by preservation bias (e.g., due to preservation/dominance of idiosomic species as you state)?
L323: I’m not convinced it makes sense to compare the testate amoeba data with the DWT, since the DWT is derived from the testate communities. A better comparison might be to the plant macrofossils. Also, the NMDS is underused, it’s not really integrated into the discussion. You could either expand the interpretation or consider removing it altogether.
L344–350: Not clear what this NMDS ordination really adds. Fire is barely discussed in the text, yet macrocharcoal plays a role in this plot. Consider dropping or restructuring this part unless you can tie it in more meaningfully. The discussion of species clustering is also underdeveloped, and could merit some statistical analysis and inclusion in the discussion.
L352–355: The note on hummock species surviving fire is interesting, maybe bring that up earlier in the discussion? Fire is not really a central theme of this paper but there are some macrocharcoal peaks.
Fig. 7: Would it be helpful to expand on how the wetness changes relate to known RCC phases? You could also consider adding the lake Żabieniec record directly into the figure for comparison?
Fig. 7 caption: Clarify—by “disturbances” do you mean anthropogenic or climatic?
L397–400: This might tie in with the chironomid-based reconstructions, could be worth pointing that out. Are we seeing cooling or drying reflected here?
Pg. 17, Discussion: I think you could say more about the implications of this work for future climate change. What can we learn from these past dynamics?
L401: It looks like there’s a phase of terrestrialisation here, possibly linked to the 8.2 ka event? It’s worth noting that this event was quite drawn-out in some records, with a lead-up to the main climate shift.
L404–405: What’s your take on the discrepancy here? Worth expanding.
L428: The change in the TA record seems to be driven by increasing A. seminulum and A. militaris—you could discuss their ecology, especially A. militaris as a dry indicator.
L439: What might have caused the disturbance observed here?
L489: E. vaginatum is a peat-former, so its presence doesn’t necessarily indicate an interruption in peat accumulation, maybe rephrase.
L506: Could you explain how the process you're referring to works, for readers less familiar with it?
L517: I’d be cautious about suggesting testate amoebae are simply better indicators than plants. They’re strongly influenced by vegetation themselves. Maybe include a bit of reflection on how this affects your interpretation.
L520–521: Could be a good place to link back to the Swindles et al. (2019) Nature Geoscience paper on the drying of European peatlands. Your data seem to support it.
L256: Linje is a well-studied site—do you know what the current average WTD is? Might be worth including here to support the reconstruction or provide some sense of its limitations.
Section 5.4: This section is quite descriptive and doesn’t fully interpret the environmental changes discussed earlier. It would be better to merge it with the previous section into a single, interpretive narrative.
L550–551: This sounds like an interesting record, maybe include it more centrally in your interpretation or even in Fig. 7?
L557–558: If this is a different core but shows synchronicity, that could support the representativeness of your record, at least for the past 2000 years.
L570: Could climate and hydrology be linked here? Or perhaps trophic shifts or mineral input played a role too- consider other possible drivers.
Technical comments
L31: Change “continue to develop” to “have continued to develop” or similar for grammatical accuracy.
L33: Insert “climatic” between “global” and “events.” Remove “(and references therein)” and instead use ‘citations from’: “cf. Birks and Seppä, 2010.”
L41: Replace “wetness” with the more precise term “bog surface wetness.”
L48: Remove the hyphen in “Mid- to Late Holocene.”
L50: “Event” should be plural, change to “events.”
L51: Replace “studied” with “identified.”
L52–55: Revise for clarity. Something like: “While there are examples of full postglacial records from Polish lakes, the longest continuous peat records from Poland come from Stążki and Głęboczek peatlands, which span 7500 and 6500 years, respectively.”
L55: Change “cores” to “sequences” or “records.”
L58: Insert “the” between “covers” and “past”; change “were conducted” to “have been conducted.”
L59: The sentence on Noryśkiewicz (2005) seems disjointed—consider moving it to join the palaeoecological section at L63–67.
L61: Change “With a” to “Generating a”; consider defining “phytocoenoses.”
L83: Replace “heritage” with “legacy,” if that’s the intended meaning.
Figure 1: Ensure all sites referenced in the text (e.g., Bydgoszcz) are shown on the map. A satellite image may be more informative than the current LiDAR-style image unless you're using it to illustrate topography/microclimates.
Fig. 1 Caption: Correct citation to “Marcisz et al., (2015).” Some minor errors like this persist throughout the paper.
L91: Replace “with a” with “using a.” Add “style” after “Russian.”
L101: Rephrase to: “low-resolution palynological data (data not published).”
L105: If this is unpublished data, consider moving it to the results rather than methods.
L108: Replace “fragment” with “section.”
L109: Rephrase: “testate amoebae were not present in significant numbers”.
L110: Replace “with tap water” with “using tap water.”
L103: Reword: “vegetation characteristic of the ombrotrophic section of the site.”
L104: Clarify which monoliths are being referenced.
L127: Add “a” to read “under a 300 µm mesh.”
L148: Correct to “Pan-European.”
Figure 3: Ensure zone 1 begins at the surface as is normal convention.
Figure Captions (all): Provide more details, including colour coding, age/depth references, and explanation of graph features. Figures should stand alone separate from the main text.
L149: Change “was also visualised as” to “were also visualised as.”
L155: Consider omitting citations to ggplot2 and ggrepel unless required by journal policy, as they weren’t actively part of any analysis.
L167: Replace “should reflect” with “likely reflects” or “should be interpreted with caution.”
L176: Suggested rephrasing: “The DWT curve illustrates the result of the testate amoeba-derived reconstruction.”
Figure 4: Improve readability—consider splitting the figure or enlarging the text, or both.
Figure 5: Remove or simplify exaggeration curves to reduce clutter. Make zone lines dashed for consistency with other figures. Increase text size for axis labels, ages, and depth units.
L184: Change “timeline” to “record.”
L192: Use “associated with” instead of “inhabit.”
L193: Clarify the meaning of “good”, consider rephrasing for specificity.
L220: Rephrase “shows that this zone is wet” to “remains wet” or “inundated.” Quantify if possible.
L222: Change “drop” to “a reduction.”
L223: Reword to something like: “…indica, the latter two species being associated with drier conditions.”
L233: Change to “increase and remain abundant until…”
L269: Replace “missing” with “absent from the record.” As they may have been present at this time, but were’nt detected in the slide.
L273: Rephrase: “contain aboveground epidermis.”
L276: Use “around” instead of “at” to reflect age uncertainty.
L283: Replace “also occurred” with “were present.”
L291: Change “when” to “indicated by.”
L293: Use “moss record” or “bryophyte record.”
L294: Change “time sequence” to “period.”
L301: Sentence is awkward—suggest rephrasing for clarity.
L315: Rephrase “appear in samples abundantly” to “are abundant.”
L324: Round R² and p-values to 2–3 decimal places and group in a single bracket. Replace “arrow” with “vector.”
Figure 6: Use a white background to enhance contrast of coloured points.
Figure 6 Caption: Report stress values in the main text.
Figure 7 Caption: Clarify what dashed lines and grey bars indicate. Why are the zones centred in the figure?
L387: Remove the extra “the” from “as these the previous dates…”
L388: Replace “encumbered” with “affected.”
L400: The chronology of events described in this section here are disordered, consider rephrasing for clearer structure.
L440: Replace “agrees” with “coincides.”
L453: For consistency, refer to this as “the 4.2 ka event.” Here and elsewhere.
L473: Replace “analysis” with “records.”
L480: Add “Sphagnum composition in this section of the record.”
L504: Remove the word “that.”
L510: Replace “is visible with a” with “occurs with a.”
L513: Rephrase to “is a very common plant on the site in the present day.
L523: Change “common” to “commonly used”; add “to reconstruct hydrological variability throughout the Holocene.”
L524: Add: “in the region throughout the past c. 11.5 ka.”
L525: Rephrase to “remarkably stable throughout the Holocene.”
L527: Replace “study timeline” with “record”; add “at the site” after “testate amoeba species.”
L553: Use ages instead of depths. Change “has also been” to “was also.”
L568: Delete “exclusively” unless there's specific evidence for full ombrotrophy.
Citation: https://doi.org/10.5194/egusphere-2025-2087-RC1 -
AC2: 'Reply on RC1', Eliise Poolma, 22 Aug 2025
We would like to sincerely thank the reviewer for the very helpful comments and suggestions that have significantly improved the quality and clarity of our manuscript. We have carefully considered all points raised and revised the text accordingly. In the specific replies below, we address each of the comments. We have expanded several explanations, improved clarity and language, and updated the reference list with more recent publications (including the references suggested by the reviewer). Regarding the NMDS ordination, we acknowledge the reviewer’s concerns, and we intend to revise this analysis accordingly (details provided in our specific replies). All technical corrections suggested by the reviewer have been implemented directly in the manuscript.
L35–38: Clarify what is meant by "main peat-forming species"; consider specifying taxa but also highlight that Sphagnum is not a prerequisite for peat formation”
Thank you for this comment. This section is clarified as follows: “Among the various vascular plants and bryophytes that contribute to peat formation, Sphagnum mosses stand out as particularly valuable in situ biological proxies in paleoecological research (Lamentowicz et al., 2019b; Ronkainen et al., 2014; Rydin et al., 2006; Clymo, 1984).”
L38: Include the citation: “https://doi.org/10.1016/0169-5347(95)90007-1” to support the ecological role of Sphagnum here. It might be helpful to add a few lines summarising its key traits.
Reference included and key traits are summarised in the following way: “The unique morphological, physiological, and biochemical traits of Sphagnum mosses make them effective ecosystem engineers, capable of building peat from their dead tissue while creating acidic, nutrient-poor, and waterlogged conditions. In doing so, they suppress the growth of vascular plants, further reinforcing conditions that favour their own dominance and continued growth (van Breemen, 1995; Turetsky et al., 2025).”
L34–36: Add a few lines discussing the value of Sphagnum in palaeoecology—it has many unique properties that make it valuable (e.g., decay resistance, vertical growth), but it's not essential for peat to form.
Regarding the two previous comments. We included the citation, emphasised that Sphagnum is not the only peat former, and added a section of key traits of Sphagnum as follows: “Among the various vascular plants and bryophytes that contribute to peat formation, Sphagnum mosses stand out as particularly valuable in situ biological proxies in paleoecological research (Lamentowicz et al., 2019b; Ronkainen et al., 2014; Rydin et al., 2006; Clymo, 1984). This is due to their distinct ecological preferences, physiological limitations, slow vertical growth, and resistance to decay. Different Sphagnum species are confined to specific microhabitats that vary in pH, nutrient availability, and water table depth (Rydin et al., 2006). Species turnover is strongly linked to changes in peatland hydrology, as Sphagnum species differ in their sensitivity to moisture availability in the capitulum and cannot actively regulate water uptake or prevent water loss (Gong et al., 2020). Together, these traits make Sphagnum mosses reliable indicators of past environmental conditions, particularly fluctuations in water table levels. In the Northern Hemisphere, Sphagnum mosses are major contributors to peat accumulation, especially in ombrotrophic bogs and poor fens where they typically dominate (Halsey et al., 2000). In contrast, peat formation in rich fens is primarily driven by brown mosses (mainly the Amblystegiaceae family), and by the roots and rhizomes of vascular plants (Jaszczuk et al., 2024). The unique morphological, physiological, and biochemical traits of Sphagnum mosses make them effective ecosystem engineers, capable of building peat from their dead tissue while creating acidic, nutrient-poor, and waterlogged conditions. In doing so, they suppress the growth of vascular plants, further reinforcing conditions that favour their own dominance and continued growth (van Breemen, 1995; Turetsky et al., 2025). Sphagnum communities growing on sufficiently deep peat deposits are highly resistant to water table changes, retaining their capacity to store carbon and continue growth even during drought conditions (Lamentowicz et al., 2019b; Marcisz et al., 2020a; Moore et al., 2021).”
L42–44: The claim that testate amoebae respond more quickly to water table changes than plants is questionable. There's an issue of circularity: vegetation and amoebae (from the same core) are not truly independent- vegetation can affect testate amoeba communities. Also, at the timescales typical of palaeoecological analysis, the resolution is often too coarse to capture these response-time differences (e.g., a 1 cm slice may represent decades or even a century or more).
It is true that testate amoebae have a dependency on vegetation. However, testate amoebae need much less time to change the community than plants. For example, van der Knaap et al. (2011) by sampling the peat in 2 mm slices that represented annual resolution, show that, while Sphagnum fuscum dominated, many different shifts in the testate amoebae community occurred .
van der Knaap, W.O., Lamentowicz, M., van Leeuwen, J.F.N., Hangartner, S., Leuenberger, M., Mauquoy, D., Goslar, T., Mitchell, E.A.D., Lamentowicz, Ł., Kamenik, C., 2011. A multi-proxy, high-resolution record of peatland development and its drivers during the last millennium from the subalpine Swiss Alps. Quaternary Science Reviews 30, 3467–3480. https://doi.org/10.1016/j.quascirev.2011.06.017
We changed the sentence in the manuscript in the following way: “While testate amoebae and plants are ecologically interconnected, testate amoebae communities can adjust species composition over shorter timescales due to their short life cycles and high turnover rates, whereas mosses and many vascular plants tend to respond more gradually to hydrological changes (Marcisz et al., 2014, Väliranta et al., 2012)”
L56: Be more specific about how anthropogenic land use change has damaged Holocene peat records, e.g., drainage, cutting, cultivation.
This section has been specified: “Full Holocene peat cores from Europe are rare because of the temporal variation in peatland succession (Ruppel et al., 2013; Stivrins, 2025) as well as drainage-based economic use ( Finlayson and Spiers, 1999; United Nations Environment Programme, 2022; Verhoeven, 2014).”
L69–72: It would help to briefly explain why this site is important for studying climate response. Who will this be useful for? You could add something like: “to better understand past ecosystem responses to climate change.”
Thank you for pointing this out. Specification added as suggested.
L94: There's missing information here: where were the radiocarbon dates processed? What material was dated? What pre-treatment methods were used? Are these your own dates or from an earlier study? If the former they should be in the results, if the latter a citation is needed.
All radiocarbon dates presented in this study are original and were obtained for this research. The dated material and laboratory codes are listed in Table 1, which we kept in the Materials and Methods, as the age-depth model based on these dates follows in the Results. All samples were processed at the Poznań Radiocarbon Laboratory (Poland) following their standard pre-treatment procedures for plant macrofossil material. As this is a well-established and routinely applied protocol in radiocarbon dating, we did not provide further methodological detail in the text.
L107: Why did you use both 5 and 10 cm³ samples? Which depths used which volume?
We used samples of varying volume because the macrofossil study had two goals: the primary one was analysing plant macrofossils, originally planned at a 10 cm resolution, and the secondary goal was to recover as many subfossil Betula nana leaves as possible for a separate study. To capture finer-scale changes of interest, especially in key sections of the profile, we increased the resolution to every 5 cm. The step of the samples (either 5 cm or 10 cm) is visible in the supplementary dataset. The studied volume will also be added to the table.
L110–135: More methodological detail needed for the testate amoeba work: what magnifications were used? Which identification guides? What was the minimum test count? And just to clarify, was a stereomicroscope also used for this part? I would usually use a high-powered microscope only.
All these information have already been included in the original manuscript – see preprint pdf, page 5 lines 125-134. Only a light microscope has been used for the analysis.
L148: It would be useful to include some detail on the transfer function model used: what type of model, what is the R² and RMSE, and maybe clarify that it was run using Rioja (and cite it).
The information about the transfer function model has already been included in the original manuscript (see the section explaining statistical methods). The R2 and RMSE of the model are explained in detail in the cited publication - Amesbury et al. 2016.
L160: The structure is a bit confusing-it feels like you have two interpretation sections: one here and one later. I’d consider merging these two sections into a single “results and interpretation” section, where each environmental change is described and interpreted in turn. This would make your paper much clearer.
We thank the reviewer for this suggestion. Our intention was to first present proxy-related changes in detail with ecological interpretation and then place the changes into a broader climate-related discussion with comparison to other studies in the next chapter. Thus we would like to keep the structure, but we have also carefully reviewed and revised the Results section so that no climatological interpretation would appear there.
L164: I’d rephrase “should reflect” as something like “we lack the chronological control to detect short-term differences in accumulation rates.” Also, instead of “treated with care,” maybe say “interpreted with caution due to this limitation.” The interpretation you give later seems reasonable, so no major concern there.
Rephrased according to the recommendation: “For the topmost 100 cm we lack the chronological control to detect short-term differences in accumulation rates. This section should thus be interpreted with caution due to this limitation, as it is based on one date and it does not reflect an exponential depth-age relationship as it should have with well-developed acrotelm and catotelm layers (Marcisz et al., 2015).”
L175: Rather than just referring the reader to the figures, I’d expand a little here. Which species are dominant? Any obvious community shifts or interpretive points worth flagging?
Thank you for this suggestion. The major changes in the proxies are outlined in the beginning of the section 5.4 (“Interpreting the proxy indicated changes”). However, we agree that moving this information where the reviewer suggests might increase clarity and flow. This will be revised in the manuscript.
L189: “Indicate the presence of a freshwater habitat” – in a peat-forming system that’s a given. Do you mean more like inundated conditions or open water? Could it have been a shallow lake phase?
Thank you for pointing this out - we did mean inundated conditions or a shallow water habitat. We also clarified this in the manuscript.
L198–199: Was the core sampled all the way to the base of the peat? Some interpretation of the factors that led to peat initiation might be interesting here if you can offer some explanation.
The core was sampled until the base of the peat, but we decided to handle the earliest stages this in a separate publication. These samples have not been analysed yet for plant macrofossils, thus we limit the discussion in this paper about the peat initiation.
L204: Replace “in the region” with “in the wider landscape” - “region” can imply a much broader area.
Replacement is made here and elsewhere.
L237: Define what is meant by “intermediate microform.”
By intermediate microform, we mean either a high lawn or low hummock microform, intermediate in the wet-dry gradient. S. medium has been found growing in hollows, lawns, and it can also extend to the hummock microform.
The section has been altered for clarity “Previously dominating wet-habitat Sphagnum subgen. Cuspidata is replaced by S. divinum/medium, which might suggest development of a drier microform (Blackford, 2000; Hassel et al., 2018).”
L261: G. discoides can indicate disturbance, yes—but it’s also found in quite stable systems. Maybe rephrase to something like: “can indicate fluctuating water tables, sometimes linked to disturbance.”
Thank you! The section is rephrased according to the suggestion
L287–288: What’s the significance of the D. rotundifolia seeds here? If this is their first appearance, could it reflect increasing nutrient deficiency?
The first occurrence of Drosera seeds is recorded in the sample dated to approximately 10,900 cal. BP. This likely reflects an increase in nutrient deficiency, as both D. rotundifolia and D. anglica, despite their slightly differing habitat preferences, are indicative of oligotrophic conditions. However, in the section spanning 4,000 to 600 cal. BP, I would not add any specific ecological interpretation to their presence, and thus this sentence is removed.
To the first appearance, we added to following section: “Differentiating between the seeds of the two Drosera species is difficult. D. rotundifolia tends to indicate lower pH and exhibits a broad ecological tolerance, whereas D. anglica is typically associated with wetter conditions and slightly higher pH levels (Banaś et al., 2023). Nonetheless, the presence of either insectivorous species implies increasing nutrient limitation in the peatland environment (Mauquoy and van Geel, 2007).”
L295: Have you got any evidence or references supporting this as an indicator of hydrological variability? What might be driving the pattern?
This interpretation was based on contrasting observation that included the co-occurrence of species indicative of both wetter and drier conditions, along with a decline in H. papilio and Sphagnum abundances. However, due to the lack of supporting references, this section has been revised as follows:
„Around 2,000 cal. BP, drier conditions are inferred from an increase in B. indica and A. seminulum abundances, accompanied by a decline in H. papilio. This shift corresponds to a reconstructed water table drop to approximately 12 cm. The timing also coincides with low Sphagnum percentages and an increase in decomposed organic matter, collectively indicating increased hydrological instability.”
L292: There seems to be a mismatch between dry-indicator testate species and wetter Sphagnum types - could this be a taphonomic/preservation issue or something else?
Around 2,000 cal. BP, the presence of moderately wet Sphagnum species contrasts with testate amoeba indicators, suggesting drier conditions. However, the overall Sphagnum percentage is low, accompanied by increased amounts of decomposed organic matter and a rise in Ericaceous remains. These patterns likely reflect a relatively rapid drying event to which the Sphagnum community was unable to respond immediately. As Sphagnum cover later recovers, a compositional shift toward slightly drier conditions is evident, marked by the replacement of Sphagnum sect. Cuspidata with S. divinum/medium.
L319: Do you think the uppermost WTD reconstructions are affected by preservation bias (e.g., due to preservation/dominance of idiosomic species as you state)?
No, we do not think that the preservation bias is an issue here. We observed often in the previous studies that presence of idiosomic species in the cores is related to landscape opening and deforestations by local communities (see e.g., Lamentowicz et al. 2013 Journal of Quaternary Science; Gałka et al. 2014 Quaternary International; Marcisz et al. 2025 Scientific Reports; Lamentowicz et al. 2020 Scientific Reports; Lamentowicz et al. 2019 Quaternary Science Reviews). Moreover, the hypothesis claiming the decomposition of idiosomic shells in peat profiles is based on a single experimental study from almost two decades ago (Swindles and Roe 2007 Paleo3) and it has not been confirmed by any other study. Therefore, we think it is speculative.
L323: I’m not convinced it makes sense to compare the testate amoeba data with the DWT, since the DWT is derived from the testate communities. A better comparison might be to the plant macrofossils. Also, the NMDS is underused, it’s not really integrated into the discussion. You could either expand the interpretation or consider removing it altogether.
Thank you for addressing this important issue. We agree that the ordination is currently underrepresented in the text. We also consider moving it to the supplementary materials. To avoid the problem of circularity in comparing testate amoebae with DWT, we will revise the NMDS by removing the DWT variable and focusing instead on the relationships between the biological proxies.
L344–350: Not clear what this NMDS ordination really adds. Fire is barely discussed in the text, yet macrocharcoal plays a role in this plot. Consider dropping or restructuring this part unless you can tie it in more meaningfully. The discussion of species clustering is also underdeveloped, and could merit some statistical analysis and inclusion in the discussion.
Thank you! We will also remove the macrocharcoal variable from the NMDS ordination and expand the discussion about the species clustering together on the plot.
L352–355: The note on hummock species surviving fire is interesting, maybe bring that up earlier in the discussion? Fire is not really a central theme of this paper but there are some macrocharcoal peaks.
Thank you for the suggestion. As mentioned before by the reviewer, fire plays a minor role in this paper and our dataset, thus we have decided to omit this part from the discussion.
Fig. 7: Would it be helpful to expand on how the wetness changes relate to known RCC phases? You could also consider adding the lake Żabieniec record directly into the figure for comparison?
Thank you for this suggestion. While we agree that linking wetness changes to known Rapid Climate Change (RCC) phases could be informative, we note that RCC intervals are generally long and characterized by fluctuating climate conditions rather than clearly defined wet or dry events. In our record, this is expressed more through phases of species turnover and hydrological instability/disturbances rather than straightforward wetness shifts. For this reason, we consider it more appropriate to expand on these connections in the main text rather than directly in the figure (where the reconstructed wetness changes from our proxies are already presented). We will also make this point more explicit in the text. As recommended, we will add the Lake Żabieniec record to Figure 7.
Fig. 7 caption: Clarify—by “disturbances” do you mean anthropogenic or climatic?
Mostly climatic but since a clear anthropogenic disturbance is visible, we will include both forms of disturbance.
L397–400: This might tie in with the chironomid-based reconstructions, could be worth pointing that out. Are we seeing cooling or drying reflected here?
Thank you for pointing this out. We do mention the cooling indicated by the chironomid record later in this paragraph (lines 404-405), but we agree that this section could benefit from some restructuring to improve clarity. The Linje record suggests either a dry event or at least a period with water table fluctuations. However, based solely on our record, it is not possible to determine whether the decomposition and bryological shift were driven by cooling or warming. Other regional records, such as varves (Zander et al., 2024) and chironomids (Kotrys et al., 2020) show contrasting results during this period.
Pg. 17, Discussion: I think you could say more about the implications of this work for future climate change. What can we learn from these past dynamics?
Thank you for this valuable suggestion. We agree that our study can be better placed in the context of future climate change. Our record shows that for millennia, the peatland was resilient to natural climate variability, maintaining peat accumulation despite repeated hydrological and vegetational shifts. However, following drainage in recent centuries, the system has not shown recovery, which highlights the limits of resilience once strong anthropogenic pressures are introduced. We will expand this in the “Conclusions”.
L401: It looks like there’s a phase of terrestrialisation here, possibly linked to the 8.2 ka event? It’s worth noting that this event was quite drawn-out in some records, with a lead-up to the main climate shift.
Thank you for this comment. We interpret the changes around this section as a transition from rich fen to poor fen and we agree that after the disturbance between 8500-8300 there is a shift to drier conditions. We also acknowledge the possibility that these changes could be linked to broader hydrological variability around the 8.2 ka event, but given the resolution and nature of our data, we prefer to remain cautious in attributing the observed changes directly to the 8.2 ka event.
L404–405: What’s your take on the discrepancy here? Worth expanding.
As mentioned before - varve record (Zander et al., 2024) and chironomids (Kotrys et al., 2020) show contrasting results during this period. Our samples show high decomposition which is followed by a Sphagnum community shift to drier microform species (while the testate amoeba record does not show significant water table changes). Since dryness and decomposition could result for example from cold, more continental winters or dry and hot summers, then this remains to be clarified in future studies. There could be some seasonal variation that our resolution does not capture.
L428: The change in the TA record seems to be driven by increasing A. seminulum and A. militaris—you could discuss their ecology, especially A. militaris as a dry indicator.
A. militaris and A. seminulum are both typically present in Sphagnum-dominated peatlands – from bogs to poor-fens, and they tend to prefer drier conditions, water tables lower than 10 cm. Alabasta militaris is often present when water tables depth is somewhere between 10 to 20 cm. All these species are indicative of drier phases on the peatland and are commonly present/well preserved in peat cores.
L439: What might have caused the disturbance observed here?
The disturbance and species turnover in Sphagnum is likely caused by Mid-Holocene cooling like on many other peatlands in Europe. We clarify this in the manuscript.
L489: E. vaginatum is a peat-former, so its presence doesn’t necessarily indicate an interruption in peat accumulation, maybe rephrase.
Thank you for the comment. We rephrased as follows: “These sites display high water table fluctuations, growth of Eriophorum vaginatum and the disappearance of S. fuscum.”
L506: Could you explain how the process you're referring to works, for readers less familiar with it?
As mentioned before, we would like to pay less attention to fire activity in the revised manuscript version as it was not of importance for the development of Linje peatland, thus we will remove this statement from the revised manuscript and only point out that the hydrological shifts might be caused by increased human activity in the region.
L517: I’d be cautious about suggesting testate amoebae are simply better indicators than plants. They’re strongly influenced by vegetation themselves. Maybe include a bit of reflection on how this affects your interpretation.
It was not our intention to suggest that testate amoebae are better indicators than plants, as we compared only the differences in reaction speed. However, to avoid a misunderstanding, we removed the problematic phrase from the revised manuscript.
L520–521: Could be a good place to link back to the Swindles et al. (2019) Nature Geoscience paper on the drying of European peatlands. Your data seem to support it.
Thank you for the suggestion, we will include this reference and link it to the text.
L256: Linje is a well-studied site—do you know what the current average WTD is? Might be worth including here to support the reconstruction or provide some sense of its limitations.
The monitoring of Linje mire is ongoing. The recent analysis published by Słowinska et al. (2022 J of Biometeorology) show that water table changes are variable in different parts of the site. Especially the north-eastern part of the site is particularly vulnerable to drought as it lays on inland dune that effectively infiltrates the surface waters. Knowing the seasonal changes of hydrology of the site we cored in the most hydrologically stable central part of Linje to obtain the most objective information.
Section 5.4: This section is quite descriptive and doesn’t fully interpret the environmental changes discussed earlier. It would be better to merge it with the previous section into a single, interpretive narrative.
Thank you for the suggestion. We agree that some parts of the Section 5.4 could be merged with other parts of the manuscript. However, we wanted to separately emphasize that the observed hydrological shifts should be interpreted with some caution as
- a) the site experiences a unique microclimate now and this could have been a case in the past
- b) observed hydrological shifts are today and could have been in the past influenced by groundwater changes.
L550–551: This sounds like an interesting record, maybe include it more centrally in your interpretation or even in Fig. 7?
We thank for this suggestion but we would like to stay with the citation and discussion because of the different data structure in the cited publication.
L557–558: If this is a different core but shows synchronicity, that could support the representativeness of your record, at least for the past 2000 years.
Thank you for pointing this out. Yes, it is a different core and it is in a rather good agreement. We will emphasize this more in the revised manuscript.
L570: Could climate and hydrology be linked here? Or perhaps trophic shifts or mineral input played a role too- consider other possible drivers.
Thank you for pointing this out. We will expand this sentence while considering other possible drivers as well.
Citation: https://doi.org/10.5194/egusphere-2025-2087-AC2
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RC2: 'Comment on egusphere-2025-2087', Anonymous Referee #2, 20 Jun 2025
Thank you for the opportunity to review the article “The most complete Holocene peat record from Central Europe: multiproxy reconstruction of postglacial wetness changes and climate events from Linje peatland, Poland” by Poolma et al.,
The paper deals with the biological proxies, i.e., bryophytes, testate amoeba, microcharcoal and radiocarbon dating of the longest peat recorded in Linje mire, Poland. The topic itself is of interest and suitable for the journal. But the paper requires some confirmations, improvements, and modifications before publication. As indicated in the attached annotated pdf file more supporting evidence is required for some of the methodologies and discussions.
General comments:
The background of this study needs to be improved, and please clarify some issues. Several studies have been conducted in the Polish peatland in general, particularly in Linje mire, as stated in the background. Therefore, I can’t see clear aims or significance compared to the previous study. If the authors attempt to test the biological proxies using bryophytes and testate amoeba, therefore emphasizing and comparing the strengths and weaknesses with other proxies (e.g., pollen, geochemical, etc) should be explained. Holocene climatic fluctuations should be elaborated more, why these events give a significant contribution to further climate predictions, robustness interpretation, or new approaches to methodology. The interconnection between Holocene climatic fluctuations (stated in the background) is not clear with the Rapid Climate Change (in discussion). Other issues are references used are relatively “old”, but somewhat the authors state that paleoclimate reconstructions using peat deposits continue to develop recently. I suggest adding new published data and references to fill and extend the research gap. I would greatly appreciate it if the authors could provide chemical data to support the interpretations related to the change in peat chemistry and nutrient availability.
The methodology section should be enhanced and improved. Samples were taken from 2019; how the samples were obtained and treated needs to be clarified. A physical description of the core is needed to figure out the samples, since no representative photos are shown. Pre-treatment of samples should be clearly stated, especially in radiocarbon dating analysis. It is well known that there are some pre-treatments using Acid-Acid or Acid-Base-Acid methods to remove some inorganic materials. Other fundamental issues that should be taken into consideration are how they distinguish the macrofossils using plant parts into specific species of plants and how they count the particles. Representative photos of macrofossils, bryophytes, and charcoal should be given to help the readers figure out the data observed and obtained. It is not clear how the authors identify specific species of bryophytes using “fragments” solely, but somehow it is interesting too. The NMDS ordination approach for statistical analysis should be reconsidered (please find detailed comments). The NMDS ordination does not support the findings, or perhaps the authors should consider the plant macrofossils parameter to be included. In addition, a lot of “peatland chemistry” records were stated without any evidence of chemical analysis!
The discussion section is interesting, but the inconsistency using specific species or parameters should be considered. For example, drier conditions in Zone-3 were characterized by the presence of Salix herbacea and Empetrum nigrum and ericacous remains; however, drier conditions in Zone-5 and 6 were indicated by Sphagnum divinum and Sphagnum sect. Acutifolia species, primarily S. rubellum and S. fuscum. I suggest adding information on dry-related species and wet-related species in Figure 4. The terminology of “disturbance” and how they define this event is also unclear. Does it reflect the drier conditions, low nutrient availability, or other factors?? Moreover, the “disturbance” events should be further explored, either anthropogenic or climatic drivers. I am not sure that nutrient availability and change in chemistry of peats can be interpreted using change in biological proxy solely; these should be strengthened by chemical analysis such as C/N ratio, sulfur content, and other biogenic elements analysis. I suggest enhancing the interpretation using the correlation between DWT to the preservation/degradation particles to find out the controlling factor on peat preservation. Lastly, since there are a lot of paleoclimate records in Linje mire (reference therein), I would recommend adding a figure that shows the results and compares them to previous records, therefore we can see clearly the significance of this study. This integration could help the readers to have a broader point of view from local-regional scales.
I suggest to add some references related to this study that might help to enhanced better interpretations.
Evans et al., (2025) https://doi.org/10.1016/j.ejop.2025.126147
Sim et al., (2023; 2021) https://doi.org/10.1016/j.quascirev.2023.108020; https://doi.org/10.1016/j.ecolind.2021.108122
Swindles et al., (2020) https://doi.org/10.1016/j.ejop.2020.125693
Swindles et al., (2019) https://www.nature.com/articles/s41561-019-0462-z
Thank you
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AC3: 'Reply on RC2', Eliise Poolma, 22 Aug 2025
We would like to thank the reviewer for the careful reading of our manuscript and for providing a detailed set of comments. We acknowledge the concerns raised regarding the background, methodological detail, and clarity of some interpretations. In the revised manuscript and in our responses, we have outlined the significance of this study more clearly in comparison to earlier work from Linje peatland and other Polish peatlands. We have also added more recent references (including those suggested by the reviewer), and updated the literature to reflect current developments in peatland palaeoecology.
In addition, we have provided more detailed explanations of the plant macrofossil and testate amoeba methodologies, as well as clarifications on core retrieval, storage, subsampling, and radiocarbon dating. We have clarified the use of the term “disturbance” and agreed with the suggestions to revise the NMDS ordination approach.
While our interpretations are based on biological proxies, we fully agree that combining these with geochemical analyses would strengthen the study. Unfortunately, such data is not available at this stage but will be considered for future research.
Attached PDF provides detailed answers to all reviewer comments.
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AC3: 'Reply on RC2', Eliise Poolma, 22 Aug 2025
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RC3: 'Comment on egusphere-2025-2087', Anonymous Referee #3, 23 Jun 2025
Comments to “The most complete Holocene peat record from Central Europe: …“ by Poolma et al.
This discussion paper presents a study investigating a Holocene record from Poland. The science is sound and the results are very interesting so that this study should certainly be published. I am not an expert on Sphagnum and testate amoeba and the comments made above by reviewer 1 and 2 are quite detailed. Therefore I only add several details which I think could be considered.
As a non-expert, I would really like a figure showing examples for the testate amoeba and bryophytes encountered.
Line 42 etc.: While I agree generally, concerning vascular plants the differences inside this group are huge (1- and 2-year cycles vs. taxa with lifespans of decades).
Line 59: The record by Noryśkiewicz (2005) covers 106 pollen samples from a neighbouring site so that it may be interesting to mention it later too or even include it (important taxa/moisture indicators?) in one of the later figures (see below). It is only mentioned here and concerning Fig. 1.
Line 90 and following: Concerning the age model, I can completely understand that sample Poz.128079 was ignored, but I am not sure if Poz-128073 was automatically excluded by the used software or by the authors – maybe in case of Poz-128073, the exclusion could be explained in a little more detail.
Linbe 97 (if not changed): …as they were…
Line 92: I guess that ‘monolith’ means a profile, but I am not sure if it is referring to the core or an extra profile. It may be okay to use the term, but I at least was a little irritated.
Line 108: ‘fragment’ could be replaced by section or a similar term. ‘Fragment’ implies something is broken.
Line 111: ‘high-power microscope’? Please explain!
Line 143: For the non-expert, a table showing the assignments might help.
Figure 2: A little bit more of the approach should be given in the figure text. It does not name used software/interpolation methods.
Figures 3-5: In the preprint, the quality of the figures is not that good. In any case, genera/species names should be in italics in all figures (as they are in 3).
Line 180 and following: The results sometimes contain already interpretations – while some readers may prefer a clearer separation and to move interpretations into the discussion, I think this is okay this way. I suggest to avoid phrases like ‘we see’ or ‘we find’, but this is also a matter of taste.
Figure 6: While this figure is interesting and important, I wondered if it could be made more easily readable. Particularly for the outer taxa, more of the names could be given, and scale numbers are very small.
Line 356 and following: As implied above, I am a little confused that the results are (well) compared with over-regional records, including pollen records, but that the pollen record by Noryśkiewicz (2005) is not mentioned here. It may even be included in Fig. 7 (arboreal pollen vs. with revised age model?).
Line 385: According to Zurek (2005).
Line 525 and following: ‘water level’ may be confusing, I assume the depth to the water table is still meant here.
Line 534 and following: Some more details could be given which kind of anthropogenic influence could cause this.
Line 590-599: I think in several sentences, a ‘,’ should be placed before the ‘and’.
Citation: https://doi.org/10.5194/egusphere-2025-2087-RC3 -
AC1: 'Reply on RC3', Eliise Poolma, 21 Aug 2025
We would like to sincerely thank the reviewer for the time and effort dedicated to carefully evaluating our manuscript. Below, we provide a detailed response to all comments.
As a non-expert, I would really like a figure showing examples for the testate amoeba and bryophytes encountered.
Thank you for your interest! The species encountered in this core are very common and already well illustrated in existing resources, which is why we chose not to include photographs in the manuscript. We would, however, have provided figures if rare taxa or potentially new fossil species had been found. For high-quality images of testate amoebae, we recommend the Arcella website (http://www.arcella.nl). For bryophytes, detailed microscope images can be found in the British Bryological Society’s species finder (https://www.britishbryologicalsociety.org.uk/learning/species-finder/). The subfossil remains look very similar to the modern specimens illustrated there.
Line 42 etc.: While I agree generally, concerning vascular plants the differences inside this group are huge (1- and 2-year cycles vs. taxa with lifespans of decades).
We appreciate this observation and agree that vascular plants show considerable variation in life history strategies, ranging from annuals to perennials and clonal species. We have updated the text accordingly:
“While testate amoebae and plants are ecologically interconnected, testate amoebae communities can adjust species composition over shorter timescales due to their short life cycles and high turnover rates, whereas mosses and perennial vascular plants tend to respond more gradually to hydrological changes (Marcisz et al., 2014; Väliranta et al., 2012).”
Line 59: The record by Noryśkiewicz (2005) covers 106 pollen samples from a neighbouring site so that it may be interesting to mention it later too or even include it (important taxa/moisture indicators?) in one of the later figures (see below). It is only mentioned here and concerning Fig. 1.
Noryśkiewicz (2005) presents only a preliminary description of pollen zones without age estimates (reported only as depths). Unfortunately, we did not find a way to meaningfully integrate these results into our figures or analyses.
Line 90 and following: Concerning the age model, I can completely understand that sample Poz.128079 was ignored, but I am not sure if Poz-128073 was automatically excluded by the used software or by the authors – maybe in case of Poz-128073, the exclusion could be explained in a little more detail.
We did preliminary calculations of the model, including all dates, Both dates Poz-128079 and Poz-128073 revealed lack of compatibility to the model. By applying each radiocarbon date to the model calculation it was not possible to produce a valid chronology. Poz-128073 was not automatically excluded but after testing models:
Resolving order: Finding R_Date Poz-128079. After excluding Poz-128079, the agreement of age-depth model (Amodel) was still equal to 0. However, both dates Poz-128074 and Poz-128073 revealed the lack of compatibility with the model (Agreement of dates to the model equal to 1%). In next step, we calculated two models X) with dates Poz-128079 and Poz-128073 excluded, and Y) with dates Poz-128079 and Poz-128074 and we compared Amodel of X) and Y). Applying the same parameters i.e P_Sequence(k0=1, interpolation=2, log10(k/k0=1) the model X revealed Amodel=66%, whereas the model Y revealed Amodel=7%. Therefore, we selected model X) as one used as a basis for absolute chronology applied for this study (profile) as it is above recommended minimum equal to 60 % (Bronk Ramsey, 2008).
To the manuscript we added a short additional explanation: “Dates Poz-128079 and Poz-128073 were distinctly younger than the neighbouring dates. Preliminary model runs, both with the full set of dates included and with only one problematic date excluded, showed that these two ages had very poor agreement with the model. Therefore, both were excluded from the final age–depth model.”
Linbe 97 (if not changed): …as they were…
Change made in the revised manuscript version
Line 92: I guess that ‘monolith’ means a profile, but I am not sure if it is referring to the core or an extra profile. It may be okay to use the term, but I at least was a little irritated.
Thank you for pointing this out. To clarify, two parallel peat profiles were originally extracted at a distance of 0.5 m. However, in this study, we analysed only one of these profiles. To avoid confusion, we have revised the text accordingly.
Line 108: ‘fragment’ could be replaced by section or a similar term. ‘Fragment’ implies something is broken.
Change made in the revised manuscript version
Line 111: ‘high-power microscope’? Please explain!
Thank you for pointing this out. We mean light microscope.
Line 143: For the non-expert, a table showing the assignments might help.
Thank you for the suggestion, we have added a table to the revised manuscript that clarifies the assignment of bryophytes to the functional group.
Figure 2: A little bit more of the approach should be given in the figure text. It does not name used software/interpolation methods.
Thank you for pointing this out, we have added the missing information to the figure caption in the revised manuscript.
Figures 3-5: In the preprint, the quality of the figures is not that good. In any case, genera/species names should be in italics in all figures (as they are in 3).
The figures quality decreased after the submission; we guess it is the case of preprint repository. We have the figures in better quality and can provide them for the final manuscript version. We will make sure all taxa names are italicised on the figures, thank you for pointing this out.
Line 180 and following: The results sometimes contain already interpretations – while some readers may prefer a clearer separation and to move interpretations into the discussion, I think this is okay this way. I suggest to avoid phrases like ‘we see’ or ‘we find’, but this is also a matter of taste.
Thank you for this comment. We acknowledge that some ecological interpretation appears in the Results section, particularly where it is closely tied to the description of proxy data. We chose this structure because we find that presenting ecological interpretation alongside the proxy changes improves clarity for the reader. In contrast, the broader climatological interpretation and comparison with other studies are in the Discussion. We have carefully reviewed the text to ensure a clearer distinction between zone descriptions and interpretation, and we have also revised the phrases mentioned.
Figure 6: While this figure is interesting and important, I wondered if it could be made more easily readable. Particularly for the outer taxa, more of the names could be given, and scale numbers are very small.
We will improve the readability of the figure as suggested
Line 356 and following: As implied above, I am a little confused that the results are (well) compared with over-regional records, including pollen records, but that the pollen record by Noryśkiewicz (2005) is not mentioned here. It may even be included in Fig. 7 (arboreal pollen vs. with revised age model?).
Thank you for this suggestion. We agree that including the study by Noryśkiewicz (2005) would indeed strengthen the regional comparison. However, as this publication only provides a general preliminary description of the pollen zones without references to age estimations (only depths), we were unfortunately not able to incorporate it further into our manuscript or figures. We will acknowledge this limitation in the revised text.
Line 385: According to Zurek (2005).
Change made in the revised manuscript version
Line 525 and following: ‘water level’ may be confusing, I assume the depth to the water table is still meant here.
Thank you for pointing this out – the phrase “water level” was replaced by “water table” or “depth to the water table” in the manuscript depending on the context.
Line 534 and following: Some more details could be given which kind of anthropogenic influence could cause this.
Thank you for pointing out this inconsistency. The Sphagnum species turnover observed 200 years ago could have both climatic (Little Ice Age) as well as anthropogenic (deforestation etc.) reasons. This is specified in the revised manuscript.
Line 590-599: I think in several sentences, a ‘,’ should be placed before the ‘and’.
Change made in the revised manuscript version
Citation: https://doi.org/10.5194/egusphere-2025-2087-AC1
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AC1: 'Reply on RC3', Eliise Poolma, 21 Aug 2025
Peer review completion
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Katarzyna Marcisz
Leeli Amon
Patryk Fiutek
Piotr Kołaczek
Karolina Leszczyńska
Dmitri Mauquoy
Michał Słowiński
Siim Veski
Friederike Wagner-Cremer
Mariusz Lamentowicz
The requested preprint has a corresponding peer-reviewed final revised paper. You are encouraged to refer to the final revised version.
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Review of The most complete Holocene peat record from Central Europe: multi-proxy reconstruction of postglacial wetness changes and climate events from Linje peatland, Poland; Eliise Poolma et al. Climate of the past
This manuscript presents a new, complete Holocene reconstruction of environmental change from Linje mire in Poland, currently the only complete peat record for this epoch in Central Europe. This alone makes this study a valuable and significant contribution, well aligned with the scope of Climate of the Past.
The title clearly reflects the contents of the manuscript, although it does not explicitly mention the anthropogenic changes observed towards the end of the record, this constitutes a relatively small (albeit significant) portion of the study and does not detract from the overall clarity or relevance of the title. The abstract provides a concise and accurate summary of the study
The study introduces new data, including a robust age-depth model based on 14 AMS radiocarbon dates, as well as stratigraphic records of botanical peat composition and testate amoebae. These are effectively used to infer hydrological variability for the site throughout the Holocene. The interpretations and conclusions are well supported by the presented data, and the study includes a thorough comparison with other palaeoecological records. However, in some instances, the manuscript could benefit from a more detailed explanation of observed changes, better contextualised within the broader peatland literature, and could benefit from some re-structuring. Doing so would further highlight the significance of this site and the value of the record. Methodologically, the study follows standard and appropriate protocols, which are clearly described and allow for reproducibility, although the NMDS ordination compares testate amoeba communities with water table reconstructions derived from the same data, which is inappropriate as obviously there will be a relationship between the two records (it is interesting that the relationship is quite weak). This analysis does not contribute much to the main discussion/findings and should be omitted or changed, perhaps to better relate plant macrofossils to the DWT reconstruction for interpretation (See specific comments). The manuscript is mostly well referenced throughout (see technical comments), with an appropriate number of citations, although several references are relatively old, and one is unpublished or still under review, this is not a major objection.
The original contributions of the authors are clearly stated. The language is generally fluent and technically accurate, though there are minor stylistic and grammatical issues in places. Suggestions have been made to improve clarity, accuracy and flow in certain sections (see technical corrections).
Specific comments
L35–38: Clarify what is meant by "main peat-forming species"; consider specifying taxa but also highlight that Sphagnum is not a prerequisite for peat formation”
L38: Include the citation: “https://doi.org/10.1016/0169-5347(95)90007-1” to support the ecological role of Sphagnum here. It might be helpful to add a few lines summarising its key traits.
L34–36: Add a few lines discussing the value of Sphagnum in palaeoecology—it has many unique properties that make it valuable (e.g., decay resistance, vertical growth), but it's not essential for peat to form.
L42–44: The claim that testate amoebae respond more quickly to water table changes than plants is questionable. There's an issue of circularity: vegetation and amoebae (from the same core) are not truly independent- vegetation can affect testate amoeba communities. Also, at the timescales typical of palaeoecological analysis, the resolution is often too coarse to capture these response-time differences (e.g., a 1 cm slice may represent decades or even a century or more).
L56: Be more specific about how anthropogenic land use change has damaged Holocene peat records, e.g., drainage, cutting, cultivation.
L69–72: It would help to briefly explain why this site is important for studying climate response. Who will this be useful for? You could add something like: “to better understand past ecosystem responses to climate change.”
L94: There's missing information here: where were the radiocarbon dates processed? What material was dated? What pre-treatment methods were used? Are these your own dates or from an earlier study? If the former they should be in the results, if the latter a citation is needed.
L107: Why did you use both 5 and 10 cm³ samples? Which depths used which volume?
L110–135: More methodological detail needed for the testate amoeba work: what magnifications were used? Which identification guides? What was the minimum test count? And just to clarify, was a stereomicroscope also used for this part? I would usually use a high-powered microscope only.
L148: It would be useful to include some detail on the transfer function model used: what type of model, what is the R² and RMSE, and maybe clarify that it was run using Rioja (and cite it).
L160: The structure is a bit confusing-it feels like you have two interpretation sections: one here and one later. I’d consider merging these two sections into a single “results and interpretation” section, where each environmental change is described and interpreted in turn. This would make your paper much clearer.
L164: I’d rephrase “should reflect” as something like “we lack the chronological control to detect short-term differences in accumulation rates.” Also, instead of “treated with care,” maybe say “interpreted with caution due to this limitation.” The interpretation you give later seems reasonable, so no major concern there.
L175: Rather than just referring the reader to the figures, I’d expand a little here. Which species are dominant? Any obvious community shifts or interpretive points worth flagging?
L189: “Indicate the presence of a freshwater habitat” – in a peat-forming system that’s a given. Do you mean more like inundated conditions or open water? Could it have been a shallow lake phase?
L198–199: Was the core sampled all the way to the base of the peat? Some interpretation of the factors that led to peat initiation might be interesting here if you can offer some explanation.
L204: Replace “in the region” with “in the wider landscape” - “region” can imply a much broader area.
L237: Define what is meant by “intermediate microform.”
L261: G. discoides can indicate disturbance, yes—but it’s also found in quite stable systems. Maybe rephrase to something like: “can indicate fluctuating water tables, sometimes linked to disturbance.”
L287–288: What’s the significance of the D. rotundifolia seeds here? If this is their first appearance, could it reflect increasing nutrient deficiency?
L295: Have you got any evidence or references supporting this as an indicator of hydrological variability? What might be driving the pattern?
L292: There seems to be a mismatch between dry-indicator testate species and wetter Sphagnum types - could this be a taphonomic/preservation issue or something else?
L319: Do you think the uppermost WTD reconstructions are affected by preservation bias (e.g., due to preservation/dominance of idiosomic species as you state)?
L323: I’m not convinced it makes sense to compare the testate amoeba data with the DWT, since the DWT is derived from the testate communities. A better comparison might be to the plant macrofossils. Also, the NMDS is underused, it’s not really integrated into the discussion. You could either expand the interpretation or consider removing it altogether.
L344–350: Not clear what this NMDS ordination really adds. Fire is barely discussed in the text, yet macrocharcoal plays a role in this plot. Consider dropping or restructuring this part unless you can tie it in more meaningfully. The discussion of species clustering is also underdeveloped, and could merit some statistical analysis and inclusion in the discussion.
L352–355: The note on hummock species surviving fire is interesting, maybe bring that up earlier in the discussion? Fire is not really a central theme of this paper but there are some macrocharcoal peaks.
Fig. 7: Would it be helpful to expand on how the wetness changes relate to known RCC phases? You could also consider adding the lake Żabieniec record directly into the figure for comparison?
Fig. 7 caption: Clarify—by “disturbances” do you mean anthropogenic or climatic?
L397–400: This might tie in with the chironomid-based reconstructions, could be worth pointing that out. Are we seeing cooling or drying reflected here?
Pg. 17, Discussion: I think you could say more about the implications of this work for future climate change. What can we learn from these past dynamics?
L401: It looks like there’s a phase of terrestrialisation here, possibly linked to the 8.2 ka event? It’s worth noting that this event was quite drawn-out in some records, with a lead-up to the main climate shift.
L404–405: What’s your take on the discrepancy here? Worth expanding.
L428: The change in the TA record seems to be driven by increasing A. seminulum and A. militaris—you could discuss their ecology, especially A. militaris as a dry indicator.
L439: What might have caused the disturbance observed here?
L489: E. vaginatum is a peat-former, so its presence doesn’t necessarily indicate an interruption in peat accumulation, maybe rephrase.
L506: Could you explain how the process you're referring to works, for readers less familiar with it?
L517: I’d be cautious about suggesting testate amoebae are simply better indicators than plants. They’re strongly influenced by vegetation themselves. Maybe include a bit of reflection on how this affects your interpretation.
L520–521: Could be a good place to link back to the Swindles et al. (2019) Nature Geoscience paper on the drying of European peatlands. Your data seem to support it.
L256: Linje is a well-studied site—do you know what the current average WTD is? Might be worth including here to support the reconstruction or provide some sense of its limitations.
Section 5.4: This section is quite descriptive and doesn’t fully interpret the environmental changes discussed earlier. It would be better to merge it with the previous section into a single, interpretive narrative.
L550–551: This sounds like an interesting record, maybe include it more centrally in your interpretation or even in Fig. 7?
L557–558: If this is a different core but shows synchronicity, that could support the representativeness of your record, at least for the past 2000 years.
L570: Could climate and hydrology be linked here? Or perhaps trophic shifts or mineral input played a role too- consider other possible drivers.
Technical comments
L31: Change “continue to develop” to “have continued to develop” or similar for grammatical accuracy.
L33: Insert “climatic” between “global” and “events.” Remove “(and references therein)” and instead use ‘citations from’: “cf. Birks and Seppä, 2010.”
L41: Replace “wetness” with the more precise term “bog surface wetness.”
L48: Remove the hyphen in “Mid- to Late Holocene.”
L50: “Event” should be plural, change to “events.”
L51: Replace “studied” with “identified.”
L52–55: Revise for clarity. Something like: “While there are examples of full postglacial records from Polish lakes, the longest continuous peat records from Poland come from Stążki and Głęboczek peatlands, which span 7500 and 6500 years, respectively.”
L55: Change “cores” to “sequences” or “records.”
L58: Insert “the” between “covers” and “past”; change “were conducted” to “have been conducted.”
L59: The sentence on Noryśkiewicz (2005) seems disjointed—consider moving it to join the palaeoecological section at L63–67.
L61: Change “With a” to “Generating a”; consider defining “phytocoenoses.”
L83: Replace “heritage” with “legacy,” if that’s the intended meaning.
Figure 1: Ensure all sites referenced in the text (e.g., Bydgoszcz) are shown on the map. A satellite image may be more informative than the current LiDAR-style image unless you're using it to illustrate topography/microclimates.
Fig. 1 Caption: Correct citation to “Marcisz et al., (2015).” Some minor errors like this persist throughout the paper.
L91: Replace “with a” with “using a.” Add “style” after “Russian.”
L101: Rephrase to: “low-resolution palynological data (data not published).”
L105: If this is unpublished data, consider moving it to the results rather than methods.
L108: Replace “fragment” with “section.”
L109: Rephrase: “testate amoebae were not present in significant numbers”.
L110: Replace “with tap water” with “using tap water.”
L103: Reword: “vegetation characteristic of the ombrotrophic section of the site.”
L104: Clarify which monoliths are being referenced.
L127: Add “a” to read “under a 300 µm mesh.”
L148: Correct to “Pan-European.”
Figure 3: Ensure zone 1 begins at the surface as is normal convention.
Figure Captions (all): Provide more details, including colour coding, age/depth references, and explanation of graph features. Figures should stand alone separate from the main text.
L149: Change “was also visualised as” to “were also visualised as.”
L155: Consider omitting citations to ggplot2 and ggrepel unless required by journal policy, as they weren’t actively part of any analysis.
L167: Replace “should reflect” with “likely reflects” or “should be interpreted with caution.”
L176: Suggested rephrasing: “The DWT curve illustrates the result of the testate amoeba-derived reconstruction.”
Figure 4: Improve readability—consider splitting the figure or enlarging the text, or both.
Figure 5: Remove or simplify exaggeration curves to reduce clutter. Make zone lines dashed for consistency with other figures. Increase text size for axis labels, ages, and depth units.
L184: Change “timeline” to “record.”
L192: Use “associated with” instead of “inhabit.”
L193: Clarify the meaning of “good”, consider rephrasing for specificity.
L220: Rephrase “shows that this zone is wet” to “remains wet” or “inundated.” Quantify if possible.
L222: Change “drop” to “a reduction.”
L223: Reword to something like: “…indica, the latter two species being associated with drier conditions.”
L233: Change to “increase and remain abundant until…”
L269: Replace “missing” with “absent from the record.” As they may have been present at this time, but were’nt detected in the slide.
L273: Rephrase: “contain aboveground epidermis.”
L276: Use “around” instead of “at” to reflect age uncertainty.
L283: Replace “also occurred” with “were present.”
L291: Change “when” to “indicated by.”
L293: Use “moss record” or “bryophyte record.”
L294: Change “time sequence” to “period.”
L301: Sentence is awkward—suggest rephrasing for clarity.
L315: Rephrase “appear in samples abundantly” to “are abundant.”
L324: Round R² and p-values to 2–3 decimal places and group in a single bracket. Replace “arrow” with “vector.”
Figure 6: Use a white background to enhance contrast of coloured points.
Figure 6 Caption: Report stress values in the main text.
Figure 7 Caption: Clarify what dashed lines and grey bars indicate. Why are the zones centred in the figure?
L387: Remove the extra “the” from “as these the previous dates…”
L388: Replace “encumbered” with “affected.”
L400: The chronology of events described in this section here are disordered, consider rephrasing for clearer structure.
L440: Replace “agrees” with “coincides.”
L453: For consistency, refer to this as “the 4.2 ka event.” Here and elsewhere.
L473: Replace “analysis” with “records.”
L480: Add “Sphagnum composition in this section of the record.”
L504: Remove the word “that.”
L510: Replace “is visible with a” with “occurs with a.”
L513: Rephrase to “is a very common plant on the site in the present day.
L523: Change “common” to “commonly used”; add “to reconstruct hydrological variability throughout the Holocene.”
L524: Add: “in the region throughout the past c. 11.5 ka.”
L525: Rephrase to “remarkably stable throughout the Holocene.”
L527: Replace “study timeline” with “record”; add “at the site” after “testate amoeba species.”
L553: Use ages instead of depths. Change “has also been” to “was also.”
L568: Delete “exclusively” unless there's specific evidence for full ombrotrophy.