Most Southern Scots Pine Populations Are Locally Adapted to Drought for Tree Height Growth
, Noelia González-Muñoz 1, Santiago C. González-Martínez 1, Ricardo Alía 2 and Marta Benito Garzón 1Round 1
Reviewer 1 Report
Overall comments:
The paper is much improved but there are crucial gaps in the background section, and I still do not understand the hypothesis. In the introduction, the authors state that height tends to show strong evidence of local adaptation despite high gene flow. Then they say that asymmetrical gene flow could reduce propensity for local adaptation but they never present evidence that there is asymmetrical gene flow in this species. Is there? Then they go on to say that this is a widely distributed species therefore populations should not express local adaptation. I am confused and I think the readers will also be confused.
The second biggest critique is that they need to clarify, again, the rationale for the hypothesis. I think the paper would benefit enormously from some discussions about vulnerability of populations at the extremes of a species’ range. There are tons of papers about this, not just for trees but other species, which would strengthen the paper both in terms of the hypothesis and broader implications.
The third biggest critique is that the authors never acknowledge the role of trait tradeoffs and how growth may not be the ultimate performance metric. Unless there are other studies that demonstrate that height growth is a key determinant of long-term persistence or population growth in this species? Especially in the discussion, there needs to be some mention of how just looking at height changes is not necessarily the full picture. It seems obvious but it’s not. And just be careful to say that you are assessing whole plant performance rather than fitness, which is much more than just height.
Minor critiques are that the broader implications need to be better discussed, because it’s not clear how the results from this study can be applied to other species (other pines, other species in Spain, other conifers). And lastly, I still found many grammatical errors and I was not able to point all of them out due to time.
Line by line edits:
Line 26: gene pools
Line 31: The basis for this hypothesis is unclear, based on the text in the abstract. It would help to clarify or rewrite to say that populations at the extremes of the species’ range might be more vulnerable to environmental shifts. Just because you are south does not mean you are at risk but there is more risk implied if you are at the very edge/boundary of where the species occurs.
Line 32: from 15-year old individuals
Line 33: where plants from 16 provenances have been planted.
Line 33: this is also confusing
Line 43: on a population’s
Line 44: what are the long-term consequences? Might there be tradeoffs?
Line 51: could increasing tree growth result in a reduction in survival or reproduction? You need to discuss tradeoffs.
Line 56: remove the word large
Line 62: this does not make sense and is not biologically correct. A species cannot express adaptation for a trait. It expresses adaptations to conditions, resulting in shifting phenotypes (traits) and genotypes. I think the authors mean to say: “despite high gene flow, populations still show evidence of local adaptation to the environment, in terms of their height.”
Line 76: allow one to
Line 86: here is where you should absolutely write about how populations at the extremes of a species range could be vulnerable. And cite a bunch of papers not just for trees but many other species. Adding this will really help to understand your rationale/hypothesis.
Line 91: and have negatively impacted tree growth
Line 93: have overlooked population-level variation with regards to adaptation and acclimation potential.
Line 96: based exclusively on species occurrence
Line 96: rewrite to remove “we”
Line 100: You said above that height tends to show evidence of local adaptation, despite high gene flow. So where is this hypothesis coming from?
Line 112: “at 15-year old measured” does not make sense grammatically
Line 159: was there a holm correction?
Line 185: “the random part was fixed” is a contradiction. Please clarify. Also change: “…and accounted for the…”
Line 187: remove starting
Line 188: also has a grammatical error but at this point, I will stop noting them. This needs more refinement.
Line 226: we spatially predicted? I think you should remove spatially and clarify in the next sentence what you mean.
Table 2: I don’t understand the rounds. Do you mean that for each model you conducted the analysis multiple times? That doesn’t make sense. I also think you need to clarify what the response variable is in the table. So for example, have “Tree height ~ global model. Tree height ~ full model minus interaction”
Table 3: the two panels should be switched
Line 291: table 3 does not show what you are referencing. I think you mean Table 4.
Line 294: there are grammatical errors in this paragraph
Figure 3: can you use a different color scheme? Maybe going from violet to green or something. So it doesn’t confuse people.
Line 329: restate your hypotheses/predictions first.
Line 342: over what time period? The temporal scale is important.
Line 370: is this speculation or has someone else studied the extent of gene flow in this species?
Line 370: is there long-distance dispersal? Short-distance dispersal would be good evidence to support limited gene flow.
Line 380: I think it’s a bit strange that the authors wait until now you discuss fitness. What about tradeoffs among traits?
Line 415: what do you mean by demography modifying expectations?
Line 417: this a very weak conclusion because it doesn’t rehash the overall aims and hypotheses, it doesn’t discuss the broader implications of the study, and it doesn’t address gaps in our knowledge or future directions. I suggest completely rewriting this part.
Line 422: why would people outside of Spain care about this? Can we apply this study to other parts of the world? Other Mediterranean regions?
Author Response
Reviewer 1
Comments and Suggestions for Authors
Overall comments:
The paper is much improved but there are crucial gaps in the background section, and I still do not understand the hypothesis. In the introduction, the authors state that height tends to show strong evidence of local adaptation despite high gene flow. Then they say that asymmetrical gene flow could reduce propensity for local adaptation but they never present evidence that there is asymmetrical gene flow in this species. Is there? Then they go on to say that this is a widely distributed species therefore populations should not express local adaptation. I am confused and I think the readers will also be confused.
The second biggest critique is that they need to clarify, again, the rationale for the hypothesis. I think the paper would benefit enormously from some discussions about vulnerability of populations at the extremes of a species’ range. There are tons of papers about this, not just for trees but other species, which would strengthen the paper both in terms of the hypothesis and broader implications.
The third biggest critique is that the authors never acknowledge the role of trait tradeoffs and how growth may not be the ultimate performance metric. Unless there are other studies that demonstrate that height growth is a key determinant of long-term persistence or population growth in this species? Especially in the discussion, there needs to be some mention of how just looking at height changes is not necessarily the full picture. It seems obvious but it’s not. And just be careful to say that you are assessing whole plant performance rather than fitness, which is much more than just height.
Response: We agree with you that without data supporting the gene flow asymmetry hypothesis, we cannot make it a central part of our introduction. Because of this we have re-written our hypothesis based upon the facts of local adaptation and the vulnerability of margin populations. The hypothesis says: “…southern Scots pine populations are locally adapted to current climate, and that expected changes in climate may lead to a decrease in tree performance”. We only mention the gene flow asymmetry in the discussion as you suggested in your previous review (see lines 377-381).
Moreover, we introduce the vulnerability of these marginal populations in the introduction (lines 91-93) and in conclusions (lines 426-428).
Finally, we put our results in a broader context by discussing the trade-offs between traits and their relationship with fitness in the discussion (lines 382-388).
Minor comments:
Minor critiques are that the broader implications need to be better discussed, because it’s not clear how the results from this study can be applied to other species (other pines, other species in Spain, other conifers). And lastly, I still found many grammatical errors and I was not able to point all of them out due to time.
Response: We highlight the implications of our findings by suggesting the importance to study southern populations case by case, as their capacity to get adapted and acclimate to the local conditions can modify a priori expectations. See lines 426-429.
Line by line edits:
Line 26: gene pools
Response: These words are absent in the new version.
Line 31: The basis for this hypothesis is unclear, based on the text in the abstract. It would help to clarify or rewrite to say that populations at the extremes of the species’ range might be more vulnerable to environmental shifts. Just because you are south does not mean you are at risk but there is more risk implied if you are at the very edge/boundary of where the species occurs.
Response: We agree with you. We have re-written this part of the abstract and have included your suggestions. We described that pines can be locally adapted. However, populations at the margins of a species’ range can be more vulnerable as they inhabit under more extreme environments. Lines 23-30.
Line 32: from 15-year old individuals
Response: Done.
Line 33: where plants from 16 provenances have been planted.
Response: Done.
Line 33: this is also confusing
Response: We have removed the word “network” which could be potentially confusing.
Line 43: on a population’s
Response: Done.
Line 44: what are the long-term consequences? Might there be tradeoffs?
Response: Conclusions have been fully re-written following your comments and we have highlighted aspects related with margin populations and their capacity to get locally adapted and acclimate to local conditions. See lines 39-44.
Line 51: could increasing tree growth result in a reduction in survival or reproduction? You need to discuss tradeoffs.
Response: We have introduced that tree mortality has increased as a consequence of more often episodes of drought and heat waves. See line 53.
Line 56: remove the word large
Response: Done.
Line 62: this does not make sense and is not biologically correct. A species cannot express adaptation for a trait. It expresses adaptations to conditions, resulting in shifting phenotypes (traits) and genotypes. I think the authors mean to say: “despite high gene flow, populations still show evidence of local adaptation to the environment, in terms of their height.”
Response: Thanks. We have corrected it.
Line 76: allow one to
Response: Done.
Line 86: here is where you should absolutely write about how populations at the extremes of a species range could be vulnerable. And cite a bunch of papers not just for trees but many other species. Adding this will really help to understand your rationale/hypothesis.
Response: Thanks. We put the focus on the study species and indicated how they are more vulnerable as they inhabit less favourable habitats where water availability constrains tree growth. See lines 91-93.
Line 91: and have negatively impacted tree growth
Response: Done.
Line 93: have overlooked population-level variation with regards to adaptation and acclimation potential.
Response: Done.
Line 96: based exclusively on species occurrence
Response: Done.
Line 96: rewrite to remove “we”
Response: Done.
Line 100: You said above that height tends to show evidence of local adaptation, despite high gene flow. So where is this hypothesis coming from?
Response: Sorry, it was a mistake. We have re-written accordingly. See lines 103-105.
Line 112: “at 15-year old measured” does not make sense grammatically
Response: Corrected.
Line 159: was there a holm correction?
Response: There was not holm correction as we do not use p-values but the strength of covariation between variables.
Line 185: “the random part was fixed” is a contradiction. Please clarify. Also change: “…and accounted for the…”
Response: Done.
Line 187: remove starting
Response: Done. Thanks.
Line 188: also has a grammatical error but at this point, I will stop noting them. This needs more refinement.
Response: We have re-written a significant part of this section to make it clearer. See lines 193-202.
Line 226: we spatially predicted? I think you should remove spatially and clarify in the next sentence what you mean.
Response: We have removed the word spatially. To make it clearer we have specified that these predictions have been done across the species’ range in Spain. See lines 240.
Table 2: I don’t understand the rounds. Do you mean that for each model you conducted the analysis multiple times? That doesn’t make sense. I also think you need to clarify what the response variable is in the table. So for example, have “Tree height ~ global model. Tree height ~ full model minus interaction”
Response: A round refers to each time a variable of the full model is assessed. For example, we started by assessing the likelihood of the two-variable interaction (round # 1), to do that we compare AIC values between the full model and the full model minus the interaction term. We have partly re-written the methodology to explain what rounds are. See lines 193-202
Table 2 has been modified following your suggestions. See lines 267-268.
Table 3: the two panels should be switched
Response: Done.
Line 291: table 3 does not show what you are referencing. I think you mean Table 4.
Response: Done. Thank you.
Line 294: there are grammatical errors in this paragraph
Response: Corrected. Thank you. See lines 304-305.
Figure 3: can you use a different color scheme? Maybe going from violet to green or something. So it doesn’t confuse people.
Response: Done.
Line 329: restate your hypotheses/predictions first.
Response: We have finally decided to start the discussion of the different findings, as it was suggested by the other referee. However, the headings of the different aspects discussed are self-explanatory making unnecessary any small introduction.
Line 342: over what time period? The temporal scale is important.
Response: Done.
Line 370: is this speculation or has someone else studied the extent of gene flow in this species?
Response: Gene flow was estimated between the same group of populations in [1]. We have cited this study in the new version.
Line 370: is there long-distance dispersal? Short-distance dispersal would be good evidence to support limited gene flow.
Response: There are low levels of gene flow among the studied populations as shown in [1] that could explain the locally adapted patterns found. This is now in the new version of the manuscript.
Line 380: I think it’s a bit strange that the authors wait until now you discuss fitness. What about tradeoffs among traits?
Response: In the new version we introduce fitness and trade-offs in the introduction and discussed them as well. See lines 68-71 and 382-388.
Line 415: what do you mean by demography modifying expectations?
Response: It was incorrect. We referred that demographic rates of this species in natural populations (not in common gardens) can be largely driven by above and below ground competition, herbivory, etc. and that impact in tree height growth and hence in future predictions. We have removed the word.
Line 417: this a very weak conclusion because it doesn’t rehash the overall aims and hypotheses, it doesn’t discuss the broader implications of the study, and it doesn’t address gaps in our knowledge or future directions. I suggest completely rewriting this part.
Response: Conclusions have been reformulated accounting for the ideas of local adaptation and margin populations. See lines 425-429.
Line 422: why would people outside of Spain care about this? Can we apply this study to other parts of the world? Other Mediterranean regions?
Response: We found that margin populations can be adapted to local conditions despite inhabiting limiting environments. This highlight the importance of analyse case by case southern populations. See lines 425-429.
Reviewer 2 Report
Authors did a serious effort. Many thanks for it. Now I recommend only some minor corrections:
Line 20: …under the RCP 8.5 scenario….
Line 39: ….concentration of RCP 8.5 scenario …..
Line 72-79: This paragraph should be seriously shortened. It is not necessary to describe a well-known principle of common gardens experiments.
Line 112: We used tree growth data of Scots….
Lines 237-241: This paragraph is not “results”. Removed this to the section “Methods”.
Line 244-245: the first sentence of the paragraph is not “results”.
Line 297: Sentences “Differences betewwen…” and “Differences in climate…” move under the table.
Lines 329-333: Redundant text. Remove it.
Lines 418-422: Conclusions is too poor. Authors should add here precise information. Why did Authors expect another result? Why results of the study are interesting for audience of international journal? Authors should add some more information about international relevance of the results.
Line 445: There are missing DOIs in section References.
Line 454: Correction of the citation is needed.
Author Response
Reviewer 2
Comments and Suggestions for Authors
Authors did a serious effort. Many thanks for it. Now I recommend only some minor corrections:
Line 20: …under the RCP 8.5 scenario….
Response: Done.
Line 39: ….concentration of RCP 8.5 scenario …..
Response: Done.
Line 72-79: This paragraph should be seriously shortened. It is not necessary to describe a well-known principle of common gardens experiments.
Response: Done.
Line 112: We used tree growth data of Scots….
Response: Done.
Lines 237-241: This paragraph is not “results”. Removed this to the section “Methods”.
Response: Done.
Line 244-245: the first sentence of the paragraph is not “results”.
Response: We have modified this sentence to make clear that the first sentence refers to the results got from the battery of linear mixed effect models fitted (see Table A2). See lines 250-252.
Line 297: Sentences “Differences betewwen…” and “Differences in climate…” move under the table.
Response: Done.
Lines 329-333: Redundant text. Remove it.
Response: Done.
Lines 418-422: Conclusions is too poor. Authors should add here precise information. Why did Authors expect another result? Why results of the study are interesting for audience of international journal? Authors should add some more information about international relevance of the results.
Response: The conclusions have been reformulated. See lines 425-429
Line 445: There are missing DOIs in section References.
Response: We have checked the references.
Line 454: Correction of the citation is needed.
Response: Done
This manuscript is a resubmission of an earlier submission. The following is a list of the peer review reports and author responses from that submission.
Round 1
Reviewer 1 Report
Overall comments: It is important to understand how past and current climate conditions interact to influence whole plant performance, and the authors have attempted to do so for various tree populations across Spain. The discussion of local adaptation versus phenotypic plasticity (starting at 2nd and 3rd paragraph of introduction) was weak. But my biggest critique is that the authors never acknowledge that the populations may not be genetically distinct. “Kawecki and Ebert 2004: Demonstrating the pattern of local adaptation in the deme × test habitat interaction for fitness requires an experiment in which samples of genotypes from the local deme and some other deme(s) are directly compared under the same environmental conditions (in the same habitat).” And of course, extending these ideas to reciprocal transplants. You can get around this by saying that you assume that the genetic composition of the local populations matches their environments. But there was no indication in the methods that these are distinct genotypes, and very little description of the experimental setup in general. This must be clarified in the introduction or discussion. It was also difficult to follow because of the grammar. I also did not see any clear hypotheses or predictions until the first figure, and I have questions about the methods (experimental design particularly, but also the mixed model structure). The figures need improvement, particularly the labels and the presentation.
Line by line edits (I did not have time to correct all of them, I suggest a native speaker look again more closely):
Line 16: Part of their natural distributions are quasi-locally
Line 17: I don’t know what punctual means. Choose a different word.
Line 19: Increasing tree height is predicted for most of the study populations with exceptions in the year 2070 following pathway RCP 8.5.
Line 21: remove plasticity
Line 22: allowing it to acclimation to variable climates.
Line 23: The impact of global climate change on tree growth and forest productivity are variable and remains difficult to predict.
Line 24: following sentence is too long
Line 28: unclear where this statement about trailing edge comes from
Line 33: 15-year old scots pine populations? Or do you mean you only used data from individuals that were 15 years old?
Line 39: from reaching an optimum height
Line 50: see edit above for line 23
Line 51: poorly worded
Line 55: however, studies of tree growth that account…
Line 64: “within a species” and remove distributions
Line 64: remove sentence about asymmetrical gene flow and move it to the discussion
Line 69-72: this is unclear. Not all suites in the northern hemisphere are water limited at their edges. Are you talking about a specific place?
Line 79-80: unclear. I don’t know what you mean.
Lines 81-83: this is poorly worded. Across the range of environmental conditions, you can observe variation within a species that results from locally adapted populations or arising from plasticity in traits. If you compared traits of a given genotype across a range of environments, you could then test for plasticity.
Line 106: not enough information here about the design. Please add more. See below.
Line 99: Either they are naturally occurring or are planted, not both
Line 108: But I really don’t understand how you chose the locations.
Line 112: were the seeds from each site mixed together? When were the seeds collected? How many plantlets were still alive? What conditions were they exposed to?
Figure 1: I don’t understand how you determined which populations were maladapted. Where does the prediction come from? I thought from your introduction that you would predict all the trailing edges were “maladapted” in the sense that they would exhibit reduced performance due to drought, compared with the intermediate range.
Line 141: If they have reduced performance at the planting site compared to the origin, you would say they exhibit strong local adaptation to the home site. I think I prefer this wording than to say they exhibit maladaptation.
Line 159: I don’t understand the thresholds. Are these the cutoff values for the correlation coefficients? If so, why only 0.08 for climate?
Line 162-3: “climate of the planting site, attributable to plastic responses, and the climate of the population origin, attributable to genetic adaptation.” You need to amend this statement. It is better if you know the genotypes are the same can you determine whether variation due to local climate is a plastic response. How do you know that the different populations (at each stand) are genetically different? Am I missing something?
Line 167: given the large number of models, and that you don’t have clear hypotheses regarding which climate traits are most important, why not use model averaging instead?
Line 176: we fixed the random part does not sound correct grammatically
Line 178: please show us the full model that you tested. I still don’t understand the structure. You have blocks and some blocks occur only within certain populations or planting sites thus they are nested. But why not have local climate be the fixed effect and climate at origin be the random effect. So you are testing whether local climate impacts the tree height conditional on the given original climate.
Line 201: does the amount of deviation matter? I wonder about a value close to zero. Would you still say maladaptive? Maybe a plant is better adapted to a particular condition (in terms of fitness) but does not experience any biologically significant fitness differences under a different condition.
Line 206: why not make this positive?
Line 214: I don’t know what this pathway means. You should offer at least a brief explanation.
Line 223: ok now I see that the threshold was for r or rho
Table 1 needs to be better presented. I was confused as to why you had a delta AIC of zero multiple times, and then I realized it was the same model. If you use lines or different shading to indicate the 4 rounds or the 4 different analyses, it would be much easier to understand.
Table 1 and 2 captions need to clarify that the response variable is tree height growth.
Figure 2, I do not understand why it is useful to show the fixed effect of climate at the population’s original site. Now that I look at this figure it does not make sense to me. I guess you could argue that their current height depends on past and current conditions (separate terms), but as I stated above, it makes more sense to me to only test for effect of current conditions given past conditions (mixed model with past as random and current as fixed). You end up generating the same figure as Figure 2 but the analytical methods are different. It ultimately is just a linear regression plot with current climate versus growth, but at different levels of past climate.
Table 3, tell us what SHM is again
Line 291: I don’t understand the use of the word punctually, and you use it many times.
Figure 3: What is DFUT-pres?
Line 302: there was a stronger effect of current environment than of home environment, which is certainly indicative of plasticity if you know the genetic makeup, but otherwise you need to explicitly state the key assumption and therefore the limitation of this study.
Line 334: then is it really maladaptive if there are little or no fitness consequences?
I'm surprised you don't discuss more of the physiological adaptations.
Reviewer 2 Report
The topic of the manuscript fits well the scope of Forests. In my opinion, the manuscript should be acceptable after major revision. Sections “Introduction”, “Methods”, and “Conclusion” need some corrections and improvements; English language seems to be very poor. For details, see my comment below.
General comments:
I think (although I am not a native English speaker) the English language in the manuscript is very poor and it needs serious corrections. Some of terms are not correct (e.g. “word-wide” in Abstract etc.).
Terminology: I think, Authors use in some cases not very typical or common terminology in the field and thus some parts of the text can be confusing or not very well for understanding (e.g. “surrounding environment” instead of “habitats” in line 24; “trailing edge” in line 28; “genetic effects” in context in line 37 despite of phenotypic plasticity can be considered as genetic effect; “tree height growth model” in line 155 etc.). Terminology in the manuscript needs to be seriously checked and improved.
Detailed comments:
Lines 14-45: Abstract is well structured, nut it needs language and terminology correction. Highlighting of main scientific findings can be shortened, but more clearly and precisely presented in the context of current knowledge-gaps. What is main novelty in findings of this study?
Line 16: What means “quasi-locally”?
Line 17: ..except punctual populations” … what it means? Is this fact important in interpretation of main findings?
Line 37: summer moisture – of what?
Line 51: Which climate zones? It must be clarified.
Line 51: What means “good enough”?
Line 53: Instead of citation [1] add here some more relevant citation by the topic from mentioned climate zones, preferably from Forests journal (e.g. doi: 10.3390/f8030082).
Line 55: Large geographical scales in this context are dealing in the study Machar et al. 2017: Biogeographic model of climate conditions for vegetation zones in Czechia.
Line 64: Avoid starting sentences using general terms as „Theoretically“.
Line 67: Remove the terms “non-local (foreign) and use appropriate (e.g. alien etc.).
Line 69: The term “trailing-edge population” needs here explanation with relevant citation. The same “maladaptation” in line 73.
Line 76-83: This paragraph is beyond the context, remove it, please.
Line 84: This statement in disputable (what about Picea abies distribution)? Clarification is needed.
Line 87: Mediterranean climate is continental?
Line 94: Distribution of the tree species on the border of geographical area does not automatically lead to vulnerability by climate change. The climate extremes are main drivers… Rephrase, please, this text.
Line 97: I miss here a brief presentation of current knowledge-gaps in topic of this study. Add this one, please, by using Web of Science.
Line 98-104: The definition of the aims of the study is not very well. Please, rework it seriously. You can start following (missing) presentation of knowledge-gaps and explain in details, what is main aim(s) of this study (in order to fill any knowledge-gaps) and explain why do you use common garden approach as appropriate methodology framework. Consider, please, in this paragraph a definition of hypotheses, which you will test in the study.
Line 109: Remove, please, this unusual citation (data available…).
Line 110-116: Please, better rephrase this paragraph and be more precise.
Line 188: It seems to be not correct capture of the figure. I cannot any predictions in the map. Improve it totally following rules of the journal.
Line 125-128: Remove this text as redundant here.
Line 129 etc: Explain precisely, please, why do you chose these mentioned variables and consider to add brief table with variables in tested (in order to shorten this paragraph).
Line 145: Explain, what means “climate normal period” or, better, avoid using this general unclear terms. The same – line 148 – what means “average climate”?
Line 155: Really, did you use the growth simulation model in the frame of terminology commonly used? Use, please, the study doi: 10.1016/j.ecoinf.2015.08.004 for better explanation of forest growth models.
Line 156: “We computed” – better should be “In order to analyse…., we used correlation analyses….” /not “correlation coefficients”…/.
Line 158: Based on exploratory analyses? What it means?
Line 158, 159: Explain, what tresholds you mean.
Line 167: AIC value – explain abbreviation if the firstly you use it.
Line 167: “bivariate mixed-effect linear model” but it needs explanation.
Line 176: This is second (following step). The process of analyses should be better explain and more logically describe, you can inspire e.g. from methodology using in the study doi: 10.3390/f9020096.
Line 192: What is „optimum …. Hopt“? Explanation is needed. Has it got specifically value for different populations?
Line 196: Please, do not use „as Wang“, but explain it here briefly.
Line 193: Avoid using “foreign population”.
Line 200-208: This your interpretation of variables analyses is very similar. Really, this can be used as a strong explanation of results? You should discuss limits and shortcomings of this approach in the section Discussion.
Line 214: Explain briefly, please, why scenario RCP 8.5.
Line 218: This sentence is not very clear. Did you visualise data?
Line 221-230: This text is not results, it is methodology. Remove it and add to the relevant part of the manuscript.
Line 230: “high percentage” – what it means? What is low? Rephrase this text.
Line 231: You say “The analysis …” but it is unclear what analyses do you mind. Add short specification.
Line 235-242: The capture of the Table 1 is not professional. The most of the capture is methodology and thus it must be removed from the capture.
Line 249: The first part of the Table 2 seems to be redundant, consider removing it.
Line 257-259: Redundant description of the figure, remove it. Better, add here interpretation of main idea of the figure presentation.
Line 263: “statistically significant” – what exactly does it means? What p-value?
Line 288-292: Here there are main findings of the study. Add, please, more detailed information about spatio-temporal predictions (areas in km2 etc, differences between current situation and 2070 – main trend?).
Lines 300-302: Redundant text, not appropriate for Discussion, remove it, please.
Lines 302-307: This text redundantly repeats summary of results, it should be moved to the section Conclusion or removed.
Line 306: See my note above, Authors should briefly discuss why they use scenario RCP 8.5.
Line 309: Say more precisely what species do you mind.
Lines 378-383: Section Conclusion is not well-written and needs serious reworking; Authors should follow rules of scientific writing for this section.
