the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Hidden behind the scene – high diversity, low connectivity of deep-sea Amphipoda in the polymetallic nodule fields in the Northeast Pacific
Abstract. The Clarion-Clipperton Zone (CCZ) situated in the central east Pacific holds major portions of manganese nodule deposits and is therefore subject to intense exploration for future deep-sea mining. However, mining rises multiple concerns. Among others about its direct or indirect impact on abyssal environment. The more, proper evaluation of deep-sea protection plans to be applied there is still hindered by insufficient knowledge of the abyssal fauna diversity and their assemblages.
Amphipoda are speciose and abundant in all marine habitats and were proven to be important food source for higher trophic levels. These crustaceans are brooders with no planktonic larval stage, so migration of species depend only on swimming capacities of adult animals. In the CCZ macrofauna-sized and relatively mobile, epibenthic Amphipoda, not collected with commonly used box corers or multicorers, remained unstudied. The present work aims to fill in this gap in knowledge by: 1) characterizing the diversity of amphipods across the CCZ and one Area of Particular Environmental Interest (APEI), 2) studying the amphipod species distribution and assemblages as well as 3) inferring the connectivity between eastern and western parts of the CCZ and other deep-sea regions.
The material was collected with epibenthic sledge from three contractor zones (UK, Singapore, Germany) and from APEI-06. Specimens were sorted into families and preferably morphospecies and subjected to cytochrome c oxidase subunit I gene (COI) barcoding.
Within total of 708 individuals 23 known and one undescribed families were identified. The barcoding allowed to obtain 581 sequences that were ascribed to 207 Molecular Operational Taxonomic Units (MOTUs). The families that dominated barcoded material in terms of abundance and species richness were: Pardaliscidae (123 ind., 44 MOTUs), Eusiridae (122 ind., 32 MOTUs) and Synopiidae (74 ind., 31 MOTUs). Only 17 known species were identified; another 48 are considered as new to science. Almost 70% of MOTUs were singletons or doubletons and 186 MOTUs were unique for the present study. The remaining 21 taxa appeared to be broadly distributed. Among the MOTUs recorded only in the CCZ majority was distributed within 400 km distance. Just six species reached or exceeded 1000 km distance of occurrence. Additionally only one species was shared between contractor areas and the studied APEI that was supposed to be preserved area representative for the mining zones. The cluster analysis of the fauna collected during three expeditions to the German contractor areas revealed almost no similarity between the West and East part of the CCZ. Additionally, within the eastern part of the studied region geographic proximity appeared to have no influence on station clustering.
The study confirmed low abundance and high species richness of deep-sea amphipod fauna of the CCZ and their highly variable assemblages. Moreover, it has shown weak connectivity between eastern and western parts of the region as well as between the contractor zones, APEIs and other deep-sea regions of the World. All the above suggest high vulnerability of the assemblages studied and, with the present state of knowledge, weak representativeness of Areas of Particular Environmental Interest for the mining zones. In order to better understand the deep-sea biodiversity and develop Regional Environmental Management Plans for the whole CCZ area two main recommendations are to be made: 1) to conduct more intensive sampling program of APEIs parallel to monitoring studies of contractor zones with the goal to better assess their usefulness as reservoirs of biodiversity for the mining areas, 2) to include the epibenthic sledge among standard gears used for sampling in order to avoid missing important part of deep-sea fauna in the monitoring studies.
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Status: closed (peer review stopped)
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RC1: 'Comment on egusphere-2025-1794', Anonymous Referee #1, 28 Jul 2025
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AC1: 'Reply on RC1', Anna M. Jażdżewska, 30 Sep 2025
Dear Reviewer,
thank you very much for your detailed work on our manuscript. We find all your comments relevant and we are sure that adressing them will greatly imporve our work.Below we will list the issues raised and provide our answers.
"I urge the authors to exercise greater caution in their conclusions regarding the effectiveness of APEI-6 as a biodiversity reservoir. While their inference aligns with previous literature, it is based on very limited sampling—only six specimens were collected from APEI-6—which restricts the strength of the conclusion."
We acknowledge the limitations of the conclusion regarding the representativeness of APEI-6 for the entire CCZ region. Following the reviewer’s suggestion, we will revise this conclusion and place greater emphasis on the need for additional sampling within the APEIs. This would enable a more robust assessment of their effectiveness as conservation areas in the event that mining activities are undertaken.
"Additionally, attention should be given to the species delimitation approach using BINs (Barcode Index Numbers). In its current form, the BIN-based delimitation appears inconsistent with the presented NJ (Neighbor-Joining) tree, and I urge the authors to carefully re-examine this analysis."
Thank you for pointing this out. The analysis itself was conducted correctly; however, an error occurred during the preparation of the figure captions. This mistake may indeed give the impression of issues with the molecular delimitation of species. In the Neighbor-Joining trees, branches were collapsed according to the BIN-based delimitation. The vertical bars labeled A1 and A2 represent the two best partitions identified by the ASAP analysis, while G indicates the GMYC analysis, and C denotes the consensus delimitation applied in the study.
"Furthermore, the manuscript would benefit from thorough language editing. There are numerous grammatical issues, such as missing or unnecessary articles and awkward sentences, that undermine the clarity of the work. More specific comments are given to the pdf attached."
Thank you for your work on our manuscript and for editing the language in the PDF file attached to your revision. After completing the revision, we will ensure that the text is reviewed by a native speaker.
On behalf of all authors,
Anna Jażdżewska
Citation: https://doi.org/10.5194/egusphere-2025-1794-AC1
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AC1: 'Reply on RC1', Anna M. Jażdżewska, 30 Sep 2025
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RC2: 'Comment on egusphere-2025-1794', Anonymous Referee #2, 09 Sep 2025
The manuscript presents valuable new molecular data on abyssal amphipod diversity in the Clarion–Clipperton Zone (CCZ), an area of high scientific and policy relevance given ongoing discussions around seabed mining. The barcoding effort is commendable, and the dataset adds to the growing baseline knowledge needed for environmental management in this region. However, several critical issues need to be addressed before the manuscript can be considered robust and suitable for publication.
Strengths
Novel dataset: The study provides an important addition to molecular baseline data, especially for non-scavenging amphipods, a group underrepresented in CCZ biodiversity research.
Policy relevance: The link to APEI (Areas of Particular Environmental Interest) representativeness is highly relevant for ISA discussions and Regional Environmental Management Plans.
Integration of methods: The parallel use of epibenthic sledge and, to a lesser extent, box corer material highlights methodological considerations important for future monitoring design.
Major Concerns
Sampling sufficiency and bias
Issues with insufficient sampling is not adequately discussed. The manuscript assumes distributional ranges and connectivity patterns based on limited collections, despite the high likelihood that most abyssal species occur at very low abundances. The risk of undersampling leading to misleading conclusions about distribution, rarity, and connectivity must be explicitly addressed and carefully qualified.
Misuse and misinterpretation of references
Miljutin et al. (2011) is inappropriately cited as relevant to DISCOL disturbances; in fact, it only addresses CCZ tracks.
Jones et al. (2017) is used as evidence of impacts, but this paper is primarily a synthesis highlighting experimental design challenges and inconclusive results in some faunal groups.
Lefaible et al. (2023, 2024) are cited as showing negative meiofaunal impacts, but results actually showed stable or increased abundances in some cases, at least in areas outside of the track.
These misrepresentations undermine the credibility of the discussion and must be corrected.
Weak referencing and unsupported statements
Several statements in the discussion and conclusions lack references or are insufficiently supported. For example, claims about species vulnerability, recolonization dynamics, and the role of small population sizes need citations (use e.g., Levins & Culver, 1971). Without stronger referencing, the arguments appear speculative.
Clarity and structure issues
Certain sections are confusing or poorly phrased, particularly the paragraph beginning with “The important role in hindering fast recovery...” which makes little sense requiring some rewriting to be comprehensible.
Similarly, the section from line 576–596 needs restructuring and proper referencing.
Treatment of number of MOTUs
Only 10 known species are mentioned out of 207 MOTUs, with 48 stated as new to science. The status of the remaining 149 MOTUs is not explained. Please clarify this.
Comparative context missing
The discussion would be strengthened by referencing comparable studies, such as Stewart et al. (2023) identifying 291 annelid species from 1177 specimens in the CCZ and discussing eg species ranges. This would contextualize the amphipod results against broader CCZ biodiversity research.
Minor Points
Some conclusions are overstated given the limited dataset, eg, claims about APEI representativeness should be framed more cautiously.
Figures and tables are useful but need clearer linkage to the key arguments in the text.
Recommendation
The manuscript has potential but currently over-interprets limited data and misuses some key references. I recommend major revision. Specifically:
Strengthen discussion of sampling limitations and potential undersampling bias.
Correct misinterpretations of disturbance and meiofauna studies.
Add missing references to support claims on rarity, extinction risk, and recolonization.
Clarify MOTU treatment and species identification.
Restructure unclear or contradictory paragraphs for clarity.
If these issues are resolved, the paper would represent a significant and timely contribution to CCZ biodiversity knowledge and to debates on deep-sea mining impacts.
Citation: https://doi.org/10.5194/egusphere-2025-1794-RC2 -
AC2: 'Reply on RC2', Anna M. Jażdżewska, 30 Sep 2025
Dear Reviewer,
thank you for your time devoted to revise our manuscript. We are happy you found our piece of work important and valuable for publication after major revision. We found your comments important and we will address them while revising the text. Below we provide answers to the specific points that were raised in your revision.
Major Concerns
"Sampling sufficiency and bias
Issues with insufficient sampling is not adequately discussed. The manuscript assumes distributional ranges and connectivity patterns based on limited collections, despite the high likelihood that most abyssal species occur at very low abundances. The risk of undersampling leading to misleading conclusions about distribution, rarity, and connectivity must be explicitly addressed and carefully qualified."
We agree that the material used in this study is limited and that abyssal species are typically found in low abundances. In line with the reviewer’s suggestion, the revised version of the manuscript will include a more thorough discussion of the challenges related to undersampling. At the same time, we wish to emphasize that the vulnerability of CCZ species also stems from their inherently low abundances.
"Misuse and misinterpretation of references
Miljutin et al. (2011) is inappropriately cited as relevant to DISCOL disturbances; in fact, it only addresses CCZ tracks.
Jones et al. (2017) is used as evidence of impacts, but this paper is primarily a synthesis highlighting experimental design challenges and inconclusive results in some faunal groups.
Lefaible et al. (2023, 2024) are cited as showing negative meiofaunal impacts, but results actually showed stable or increased abundances in some cases, at least in areas outside of the track.
These misrepresentations undermine the credibility of the discussion and must be corrected."
Thank you for your comment. In the revised version, we will ensure that the references are accurately interpreted.
"Weak referencing and unsupported statements
Several statements in the discussion and conclusions lack references or are insufficiently supported. For example, claims about species vulnerability, recolonization dynamics, and the role of small population sizes need citations (use e.g., Levins & Culver, 1971). Without stronger referencing, the arguments appear speculative.
Clarity and structure issues
Certain sections are confusing or poorly phrased, particularly the paragraph beginning with “The important role in hindering fast recovery...” which makes little sense requiring some rewriting to be comprehensible.
Similarly, the section from line 576–596 needs restructuring and proper referencing.
Treatment of number of MOTUs
Only 10 known species are mentioned out of 207 MOTUs, with 48 stated as new to science. The status of the remaining 149 MOTUs is not explained. Please clarify this.
Comparative context missing
The discussion would be strengthened by referencing comparable studies, such as Stewart et al. (2023) identifying 291 annelid species from 1177 specimens in the CCZ and discussing eg species ranges. This would contextualize the amphipod results against broader CCZ biodiversity research."
Thank you for pointing all these issues out. In the revised version, we will ensure that all relevant literature is properly cited.
Regarding the status of the delimitated MOTUs, it was possible to determine it only for a relatively small portion of the collection. Due to time constraints and the destruction of some specimens—which prevented final morphological identification but still allowed for DNA extraction—the remaining MOTUs could only be identified to higher taxonomic levels (genus or family). Further studies may reveal whether these represent known taxa or are new to science.
This will be explicitly stated in the revised version to avoid any misunderstanding. Additionally, we will ensure that the text is reviewed by a native speaker to prevent language errors that could lead to misinterpretation.Minor Points
"Some conclusions are overstated given the limited dataset, eg, claims about APEI representativeness should be framed more cautiously."
As the other reviewer also raised concerns regarding the justification of APEIs' representativeness for the CCZ, we will further revise this part of the discussion and conclusions.
"Figures and tables are useful but need clearer linkage to the key arguments in the text."
We will make sure to strengthen the link between the figures and tables and the main arguments discussed in the text.
Once more thank you for your time.
On behalf of all authors,
Anna Jażdżewska
Citation: https://doi.org/10.5194/egusphere-2025-1794-AC2
-
AC2: 'Reply on RC2', Anna M. Jażdżewska, 30 Sep 2025
Status: closed (peer review stopped)
-
RC1: 'Comment on egusphere-2025-1794', Anonymous Referee #1, 28 Jul 2025
This study represents a significant contribution to the understanding of deep-sea biodiversity in the Clarion-Clipperton Zone (CCZ), a region targeted for potential deep-sea mining. The authors focus on an often-overlooked benthic group, Amphipoda, by using material sampled primerly with an epibenthic sledge. DNA barcoding is employed to reveal an impresive amphipod diversity, including a high proportion of potentially new species as well showcase the limited connectivity between CCZ areas. The research is methodologically sound, well-structured, and clearly addresses a major gap in current knowledge of the group Amphipoda, while emphasing the value of the sampling gear (EBS).
That said, I urge the authors to exercise greater caution in their conclusions regarding the effectiveness of APEI-6 as a biodiversity reservoir. While their inference aligns with previous literature, it is based on very limited sampling—only six specimens were collected from APEI-6—which restricts the strength of the conclusion.
Additionally, attention should be given to the species delimitation approach using BINs (Barcode Index Numbers). In its current form, the BIN-based delimitation appears inconsistent with the presented NJ (Neighbor-Joining) tree, and I urge the authors to carefully re-examine this analysis. Furthermore, the manuscript would benefit from thorough language editing. There are numerous grammatical issues, such as missing or unnecessary articles and awkward sentences, that undermine the clarity of the work. More specific comments are given to the pdf attached.
For the above reasons I recommend this article for publication after its being revised, as it provides robust data that should inform both the scientific community and assist to policy decisions related to deep-sea mining in the CCZ .
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AC1: 'Reply on RC1', Anna M. Jażdżewska, 30 Sep 2025
Dear Reviewer,
thank you very much for your detailed work on our manuscript. We find all your comments relevant and we are sure that adressing them will greatly imporve our work.Below we will list the issues raised and provide our answers.
"I urge the authors to exercise greater caution in their conclusions regarding the effectiveness of APEI-6 as a biodiversity reservoir. While their inference aligns with previous literature, it is based on very limited sampling—only six specimens were collected from APEI-6—which restricts the strength of the conclusion."
We acknowledge the limitations of the conclusion regarding the representativeness of APEI-6 for the entire CCZ region. Following the reviewer’s suggestion, we will revise this conclusion and place greater emphasis on the need for additional sampling within the APEIs. This would enable a more robust assessment of their effectiveness as conservation areas in the event that mining activities are undertaken.
"Additionally, attention should be given to the species delimitation approach using BINs (Barcode Index Numbers). In its current form, the BIN-based delimitation appears inconsistent with the presented NJ (Neighbor-Joining) tree, and I urge the authors to carefully re-examine this analysis."
Thank you for pointing this out. The analysis itself was conducted correctly; however, an error occurred during the preparation of the figure captions. This mistake may indeed give the impression of issues with the molecular delimitation of species. In the Neighbor-Joining trees, branches were collapsed according to the BIN-based delimitation. The vertical bars labeled A1 and A2 represent the two best partitions identified by the ASAP analysis, while G indicates the GMYC analysis, and C denotes the consensus delimitation applied in the study.
"Furthermore, the manuscript would benefit from thorough language editing. There are numerous grammatical issues, such as missing or unnecessary articles and awkward sentences, that undermine the clarity of the work. More specific comments are given to the pdf attached."
Thank you for your work on our manuscript and for editing the language in the PDF file attached to your revision. After completing the revision, we will ensure that the text is reviewed by a native speaker.
On behalf of all authors,
Anna Jażdżewska
Citation: https://doi.org/10.5194/egusphere-2025-1794-AC1
-
AC1: 'Reply on RC1', Anna M. Jażdżewska, 30 Sep 2025
-
RC2: 'Comment on egusphere-2025-1794', Anonymous Referee #2, 09 Sep 2025
The manuscript presents valuable new molecular data on abyssal amphipod diversity in the Clarion–Clipperton Zone (CCZ), an area of high scientific and policy relevance given ongoing discussions around seabed mining. The barcoding effort is commendable, and the dataset adds to the growing baseline knowledge needed for environmental management in this region. However, several critical issues need to be addressed before the manuscript can be considered robust and suitable for publication.
Strengths
Novel dataset: The study provides an important addition to molecular baseline data, especially for non-scavenging amphipods, a group underrepresented in CCZ biodiversity research.
Policy relevance: The link to APEI (Areas of Particular Environmental Interest) representativeness is highly relevant for ISA discussions and Regional Environmental Management Plans.
Integration of methods: The parallel use of epibenthic sledge and, to a lesser extent, box corer material highlights methodological considerations important for future monitoring design.
Major Concerns
Sampling sufficiency and bias
Issues with insufficient sampling is not adequately discussed. The manuscript assumes distributional ranges and connectivity patterns based on limited collections, despite the high likelihood that most abyssal species occur at very low abundances. The risk of undersampling leading to misleading conclusions about distribution, rarity, and connectivity must be explicitly addressed and carefully qualified.
Misuse and misinterpretation of references
Miljutin et al. (2011) is inappropriately cited as relevant to DISCOL disturbances; in fact, it only addresses CCZ tracks.
Jones et al. (2017) is used as evidence of impacts, but this paper is primarily a synthesis highlighting experimental design challenges and inconclusive results in some faunal groups.
Lefaible et al. (2023, 2024) are cited as showing negative meiofaunal impacts, but results actually showed stable or increased abundances in some cases, at least in areas outside of the track.
These misrepresentations undermine the credibility of the discussion and must be corrected.
Weak referencing and unsupported statements
Several statements in the discussion and conclusions lack references or are insufficiently supported. For example, claims about species vulnerability, recolonization dynamics, and the role of small population sizes need citations (use e.g., Levins & Culver, 1971). Without stronger referencing, the arguments appear speculative.
Clarity and structure issues
Certain sections are confusing or poorly phrased, particularly the paragraph beginning with “The important role in hindering fast recovery...” which makes little sense requiring some rewriting to be comprehensible.
Similarly, the section from line 576–596 needs restructuring and proper referencing.
Treatment of number of MOTUs
Only 10 known species are mentioned out of 207 MOTUs, with 48 stated as new to science. The status of the remaining 149 MOTUs is not explained. Please clarify this.
Comparative context missing
The discussion would be strengthened by referencing comparable studies, such as Stewart et al. (2023) identifying 291 annelid species from 1177 specimens in the CCZ and discussing eg species ranges. This would contextualize the amphipod results against broader CCZ biodiversity research.
Minor Points
Some conclusions are overstated given the limited dataset, eg, claims about APEI representativeness should be framed more cautiously.
Figures and tables are useful but need clearer linkage to the key arguments in the text.
Recommendation
The manuscript has potential but currently over-interprets limited data and misuses some key references. I recommend major revision. Specifically:
Strengthen discussion of sampling limitations and potential undersampling bias.
Correct misinterpretations of disturbance and meiofauna studies.
Add missing references to support claims on rarity, extinction risk, and recolonization.
Clarify MOTU treatment and species identification.
Restructure unclear or contradictory paragraphs for clarity.
If these issues are resolved, the paper would represent a significant and timely contribution to CCZ biodiversity knowledge and to debates on deep-sea mining impacts.
Citation: https://doi.org/10.5194/egusphere-2025-1794-RC2 -
AC2: 'Reply on RC2', Anna M. Jażdżewska, 30 Sep 2025
Dear Reviewer,
thank you for your time devoted to revise our manuscript. We are happy you found our piece of work important and valuable for publication after major revision. We found your comments important and we will address them while revising the text. Below we provide answers to the specific points that were raised in your revision.
Major Concerns
"Sampling sufficiency and bias
Issues with insufficient sampling is not adequately discussed. The manuscript assumes distributional ranges and connectivity patterns based on limited collections, despite the high likelihood that most abyssal species occur at very low abundances. The risk of undersampling leading to misleading conclusions about distribution, rarity, and connectivity must be explicitly addressed and carefully qualified."
We agree that the material used in this study is limited and that abyssal species are typically found in low abundances. In line with the reviewer’s suggestion, the revised version of the manuscript will include a more thorough discussion of the challenges related to undersampling. At the same time, we wish to emphasize that the vulnerability of CCZ species also stems from their inherently low abundances.
"Misuse and misinterpretation of references
Miljutin et al. (2011) is inappropriately cited as relevant to DISCOL disturbances; in fact, it only addresses CCZ tracks.
Jones et al. (2017) is used as evidence of impacts, but this paper is primarily a synthesis highlighting experimental design challenges and inconclusive results in some faunal groups.
Lefaible et al. (2023, 2024) are cited as showing negative meiofaunal impacts, but results actually showed stable or increased abundances in some cases, at least in areas outside of the track.
These misrepresentations undermine the credibility of the discussion and must be corrected."
Thank you for your comment. In the revised version, we will ensure that the references are accurately interpreted.
"Weak referencing and unsupported statements
Several statements in the discussion and conclusions lack references or are insufficiently supported. For example, claims about species vulnerability, recolonization dynamics, and the role of small population sizes need citations (use e.g., Levins & Culver, 1971). Without stronger referencing, the arguments appear speculative.
Clarity and structure issues
Certain sections are confusing or poorly phrased, particularly the paragraph beginning with “The important role in hindering fast recovery...” which makes little sense requiring some rewriting to be comprehensible.
Similarly, the section from line 576–596 needs restructuring and proper referencing.
Treatment of number of MOTUs
Only 10 known species are mentioned out of 207 MOTUs, with 48 stated as new to science. The status of the remaining 149 MOTUs is not explained. Please clarify this.
Comparative context missing
The discussion would be strengthened by referencing comparable studies, such as Stewart et al. (2023) identifying 291 annelid species from 1177 specimens in the CCZ and discussing eg species ranges. This would contextualize the amphipod results against broader CCZ biodiversity research."
Thank you for pointing all these issues out. In the revised version, we will ensure that all relevant literature is properly cited.
Regarding the status of the delimitated MOTUs, it was possible to determine it only for a relatively small portion of the collection. Due to time constraints and the destruction of some specimens—which prevented final morphological identification but still allowed for DNA extraction—the remaining MOTUs could only be identified to higher taxonomic levels (genus or family). Further studies may reveal whether these represent known taxa or are new to science.
This will be explicitly stated in the revised version to avoid any misunderstanding. Additionally, we will ensure that the text is reviewed by a native speaker to prevent language errors that could lead to misinterpretation.Minor Points
"Some conclusions are overstated given the limited dataset, eg, claims about APEI representativeness should be framed more cautiously."
As the other reviewer also raised concerns regarding the justification of APEIs' representativeness for the CCZ, we will further revise this part of the discussion and conclusions.
"Figures and tables are useful but need clearer linkage to the key arguments in the text."
We will make sure to strengthen the link between the figures and tables and the main arguments discussed in the text.
Once more thank you for your time.
On behalf of all authors,
Anna Jażdżewska
Citation: https://doi.org/10.5194/egusphere-2025-1794-AC2
-
AC2: 'Reply on RC2', Anna M. Jażdżewska, 30 Sep 2025
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This study represents a significant contribution to the understanding of deep-sea biodiversity in the Clarion-Clipperton Zone (CCZ), a region targeted for potential deep-sea mining. The authors focus on an often-overlooked benthic group, Amphipoda, by using material sampled primerly with an epibenthic sledge. DNA barcoding is employed to reveal an impresive amphipod diversity, including a high proportion of potentially new species as well showcase the limited connectivity between CCZ areas. The research is methodologically sound, well-structured, and clearly addresses a major gap in current knowledge of the group Amphipoda, while emphasing the value of the sampling gear (EBS).
That said, I urge the authors to exercise greater caution in their conclusions regarding the effectiveness of APEI-6 as a biodiversity reservoir. While their inference aligns with previous literature, it is based on very limited sampling—only six specimens were collected from APEI-6—which restricts the strength of the conclusion.
Additionally, attention should be given to the species delimitation approach using BINs (Barcode Index Numbers). In its current form, the BIN-based delimitation appears inconsistent with the presented NJ (Neighbor-Joining) tree, and I urge the authors to carefully re-examine this analysis. Furthermore, the manuscript would benefit from thorough language editing. There are numerous grammatical issues, such as missing or unnecessary articles and awkward sentences, that undermine the clarity of the work. More specific comments are given to the pdf attached.
For the above reasons I recommend this article for publication after its being revised, as it provides robust data that should inform both the scientific community and assist to policy decisions related to deep-sea mining in the CCZ .