An 11 000-year-old giant muntjac subfossil from Northern Vietnam: implications for past and present populations

Described at the end of the twentieth century, the large-antlered or giant muntjac, Muntiacus gigas (syn. vuquangensis), is a Critically Endangered species currently restricted to the Annamite region in Southeast Asia. Here we report subfossil evidence of giant muntjac, a mandible fragment dated between 11.1 and 11.4 thousand years before present, from northern Vietnam. We describe morphological and metric criteria for diagnosis and consider the specimen in the context of regional archaeological and palaeontological records of Muntiacus. We then consider the palaeoenvironmental context of the specimen and the implications for habitat requirements for extant populations. The new specimen extends the known spatial and temporal range of giant muntjacs in Vietnam and is further evidence that this species was more widely distributed in the Holocene than current records indicate. While regional proxy evidence indicates a drier climate and more open woodland habitats at the onset of the Holocene, contextual evidence indicates that the specimen derived from an animal inhabiting limestone karst forest. This record also supports the assertion that remnant populations are in a refugial state, as a result of anthropogenic pressures, rather than representing a centre of endemism. These facts underscore the urgent need for the conservation of remaining populations.


2
Have you any concerns about statistical analyses in this paper? Yes

Recommendation?
Major revision is needed (please make suggestions in comments)

Comments to the Author(s)
The paper by Stimpson et al. provides important new historical baseline data on a Critically Endangered but largely overlooked mammal species, with potentially important implications for understanding its ecological requirements and thus informing future conservation management. However, additional analysis and some restructuring of the manuscript is required before it can be published.
Major comments: -The introduction is all a bit too specific, and makes the paper feel like it would only appeal to someone already interested in muntjacs. It would be much better to open the paper by framing the study in a much wider context, by starting off with an explanation about the importance of using data from the past (historical baselines, environmental archives) to provide new insights for conservation of threatened species, and the ways in which this can be / has been done -i.e. make the paper of interest to people beyond just muntjac specialists! -Similarly, the previous suggestion that vuquangensis and gigas are the same species should be introduced in the introduction, rather than this important point being relegated to the discussion -as your study provides further evidence of a past wider distribution of giant muntjacs, which supports the previous study.
-Similarly, the idea that the name "gigas" can be "ignored" for now, even though it represents a senior synonym of vuquangensis, because "this revision has not yet been formally adopted", is a bit of a red herring -the way it can be adopted is by studies like this deciding to use the name "gigas" (it doesn't require a formal IUCN edict) -as there is no worldwide arbiter of taxonomy. So, if the authors consider that vuquangensis and gigas are the same taxon, they should use the name gigas in this manuscript; and if not, then they should state why they think they're different.
-When comparing measurements between the new specimen and existing data for other muntjacs, the authors should also compare their specimen with available measurement data for Muntiacus gigas -this is really important to do.
-The lack of any statistical analysis to compare the new specimen with other muntjac samples (muntiacus, vuquangensis, gigas) is a major omission, and really has to be done -even if just via something fairly straightforward such as t-tests or ANOVAs. A statistical assessment of measurement differences, rather than just "eyeballing" the data, is essential.
Minor comments: -"sub-fossil" should be changed throughout the manuscript to "subfossil" (the more normal way of spelling this word) -"critically endangered" should be capitalised to "Critically Endangered", as this is a formal IUCN threat category. It also shouldn't be written in inverted commas (line 58) -Line 112: "11, 778" -spacing after the comma needs correcting 3 -Line 243: "ectosylids" spelled incorrectly -Line 257: "Muntiacini" shouldn't be italicised -Line 263: "celaphodus" spelled incorrectly -Line 285: not strictly true that the only confirmed gigas specimens derive from Holocene sites in China -Turvey et al 2016 also included a Late Pleistocene specimen from Yuhang in their analyses, so Pleistocene material has been interpreted by previous authors as representing gigas -Line 352-3: even if lower elevational estimates are confounded by human activity, it would still be useful to say what the current lower elevational range estimate is for the species, and how much this differs from the elevation of the Holocene site from which the new specimen was found -and also discuss the idea of the Annamites acting as an altitudinal refugium for the species, in the context of this concept more widely for SE Asian and other mammal species 5 Decision letter (RSOS-181461.R0) 17-Dec-2018 Dear Dr Stimpson, The editors assigned to your paper ("An 11,000-year-old giant muntjac (Muntiacus vuquangensis) sub-fossil from Northern Vietnam: implications for past and present populations") have now received comments from reviewers. We would like you to revise your paper in accordance with the referee and Associate Editor suggestions which can be found below (not including confidential reports to the Editor). Please note this decision does not guarantee eventual acceptance.
Please submit a copy of your revised paper before 09-Jan-2019. Please note that the revision deadline will expire at 00.00am on this date. If we do not hear from you within this time then it will be assumed that the paper has been withdrawn. In exceptional circumstances, extensions may be possible if agreed with the Editorial Office in advance. We do not allow multiple rounds of revision so we urge you to make every effort to fully address all of the comments at this stage. If deemed necessary by the Editors, your manuscript will be sent back to one or more of the original reviewers for assessment. If the original reviewers are not available, we may invite new reviewers.
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If you wish to submit your supporting data or code to Dryad (http://datadryad.org/), or modify your current submission to dryad, please use the following link: http://datadryad.org/submit?journalID=RSOS&manu=RSOS-181461 • Competing interests Please declare any financial or non-financial competing interests, or state that you have no competing interests. • Authors' contributions All submissions, other than those with a single author, must include an Authors' Contributions section which individually lists the specific contribution of each author. The list of Authors should meet all of the following criteria; 1) substantial contributions to conception and design, or acquisition of data, or analysis and interpretation of data; 2) drafting the article or revising it critically for important intellectual content; and 3) final approval of the version to be published.
All contributors who do not meet all of these criteria should be included in the acknowledgements.
We suggest the following format: AB carried out the molecular lab work, participated in data analysis, carried out sequence alignments, participated in the design of the study and drafted the manuscript; CD carried out the statistical analyses; EF collected field data; GH conceived of the study, designed the study, coordinated the study and helped draft the manuscript. All authors gave final approval for publication.
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Once again, thank you for submitting your manuscript to Royal Society Open Science and I look forward to receiving your revision. If you have any questions at all, please do not hesitate to get in touch. Comments to the Author(s) The paper by Stimpson et al. provides important new historical baseline data on a Critically Endangered but largely overlooked mammal species, with potentially important implications for understanding its ecological requirements and thus informing future conservation management. However, additional analysis and some restructuring of the manuscript is required before it can be published.

Major comments:
-The introduction is all a bit too specific, and makes the paper feel like it would only appeal to someone already interested in muntjacs. It would be much better to open the paper by framing the study in a much wider context, by starting off with an explanation about the importance of using data from the past (historical baselines, environmental archives) to provide new insights for conservation of threatened species, and the ways in which this can be / has been done -i.e. make the paper of interest to people beyond just muntjac specialists! -Similarly, the previous suggestion that vuquangensis and gigas are the same species should be introduced in the introduction, rather than this important point being relegated to the discussion -as your study provides further evidence of a past wider distribution of giant muntjacs, which supports the previous study.
-Similarly, the idea that the name "gigas" can be "ignored" for now, even though it represents a senior synonym of vuquangensis, because "this revision has not yet been formally adopted", is a bit of a red herring -the way it can be adopted is by studies like this deciding to use the name "gigas" (it doesn't require a formal IUCN edict) -as there is no worldwide arbiter of taxonomy. So, if the authors consider that vuquangensis and gigas are the same taxon, they should use the name gigas in this manuscript; and if not, then they should state why they think they're different.
-When comparing measurements between the new specimen and existing data for other muntjacs, the authors should also compare their specimen with available measurement data for Muntiacus gigas -this is really important to do.
-The lack of any statistical analysis to compare the new specimen with other muntjac samples (muntiacus, vuquangensis, gigas) is a major omission, and really has to be done -even if just via something fairly straightforward such as t-tests or ANOVAs. A statistical assessment of measurement differences, rather than just "eyeballing" the data, is essential.
Minor comments: -"sub-fossil" should be changed throughout the manuscript to "subfossil" (the more normal way of spelling this word) -"critically endangered" should be capitalised to "Critically Endangered", as this is a formal IUCN threat category. It also shouldn't be written in inverted commas (line 58) -Line 112: "11, 778" -spacing after the comma needs correcting -Line 243: "ectosylids" spelled incorrectly -Line 257: "Muntiacini" shouldn't be italicised -Line 263: "celaphodus" spelled incorrectly -Line 285: not strictly true that the only confirmed gigas specimens derive from Holocene sites in China -Turvey et al 2016 also included a Late Pleistocene specimen from Yuhang in their analyses, so Pleistocene material has been interpreted by previous authors as representing gigas -Line 352-3: even if lower elevational estimates are confounded by human activity, it would still be useful to say what the current lower elevational range estimate is for the species, and how much this differs from the elevation of the Holocene site from which the new specimen was found -and also discuss the idea of the Annamites acting as an altitudinal refugium for the species, in the context of this concept more widely for SE Asian and other mammal species Reviewer: 2 Comments to the Author(s) Dear authors, This is a carefully conducted transdisciplinary study, a very well written manuscript, and an excellent example that we need the big picture, far beyond historical time, when assessing conservational needs. Congratulations, I've enjoyed reading very much! I've only indicated minor suggestions to consider in the manuscript. The two most important among them, I'm coming up with here again: Critical data of Capreolus pygargus and Elaphodus cephalophus megalodon seem to have been neglected. In order to make the issues more transparent I'd suggest to add relevant data. Two issue Reviewer: 3 Comments to the Author(s) The manuscript is well written and if the topic falls into the scope of the journal, it definitely is worth to be published. Unfortunately I have not found any access to the supplementary tables and figures. Therefore, I could not comment some of the evidence used in the ms. Substantial part of the arguments is based on the body mass estimation. The method is recalled from Janis (1990). Since the reference deals with a chapter from a book published almost 30 years ago, it may be difficult for many readers to get and read the methodology there. This may have two implications to the recent manuscript. First, the authors of the present study should describe in brief the principles of the method for those readers who do not have the cited book at hand. Second, from a more recent literature (e. g., Mendoza, M., Janis, C. M. & Palmqvist, P., 2006. Estimating the body mass of extinct ungulates: a study on the use of multiple regression. J. Zool. 270, 90-101; De Esteban-Trivigno, S. & Köhler, M., 2011. New equations for body mass estimation in bovids: Testing some procedures when constructing regression functions. Mamm. Biol. 76, 755-761) it is apparent that some improvements of the method are available since 1990. Even if it was not applicable to the paleo-material available, it seems the body mass estimation based on multiple regression should be considered more carefully.
Ad Table 1

Comments to the Author(s)
The authors have done a great job in addressing all of my comments and suggestions, and this is a great paper -I look forward to it being published! The only very minor further edits that I'd suggest are a few typos here and there: -Page 3, lines 56-58: It feels like there's a word missing from this sentence -maybe it needs a "For" at the start of the sentence? -Page 5, line 47: should "Primoye" be "Primorye"? -Page 5, line 57: "temperature" should be "temperate" -Page 6, line 4: remove "and" after "biostratigraphic" -Page 6, line 50: should "(NHMUK 2010)" be removed? (not sure what this means after the actual reported specimen number) -Page 7, line 49: remove "of" from before "are present in Quaternary archives" -Page 8, line 3: remove "of" after "published metric identifications" -Page 8, line 6: "post glacial" is usually either hyphenated or one word - Figure 3: "Hydropotes" is spelled wrong in the figure, and "Capreolus (capreolus) pygargus" should have the subgenus name in parentheses removed (since it's incorrectly capitalised, and you don't refer to this subgenus name in the main text)

12-Feb-2019
Dear Dr Stimpson: On behalf of the Editors, I am pleased to inform you that your Manuscript RSOS-181461.R1 entitled "An 11,000-year-old giant muntjac subfossil from Northern Vietnam: implications for past and present populations" has been accepted for publication in Royal Society Open Science subject to minor revision in accordance with the referee suggestions. Please find the referees' comments at the end of this email.
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• Data accessibility It is a condition of publication that all supporting data are made available either as supplementary information or preferably in a suitable permanent repository. The data accessibility section should state where the article's supporting data can be accessed. This section should also include details, where possible of where to access other relevant research materials such as statistical tools, protocols, software etc can be accessed. If the data has been deposited in an external repository this section should list the database, accession number and link to the DOI for all data from the article that has been made publicly available. Data sets that have been deposited in an external repository and have a DOI should also be appropriately cited in the manuscript and included in the reference list.
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We suggest the following format: AB carried out the molecular lab work, participated in data analysis, carried out sequence alignments, participated in the design of the study and drafted the manuscript; CD carried out the statistical analyses; EF collected field data; GH conceived of the study, designed the study, coordinated the study and helped draft the manuscript. All authors gave final approval for publication.
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RESPONSE TO REVIEWERS
We thank the referees and editors for their time reviewing our submission "An 11,000-year-old giant muntjac subfossil from Northern Vietnam: implications for past and present populations". The comments and suggested revisions improve the content of the revised manuscript and the acknowledgements have been updated, accordingly. We are particularly grateful to Reviewer 1 for supplying details of the dental dimensions of Muntiacus gigas.

REVIEWER 1 The paper by Stimpson et al. provides important new historical baseline data on a Critically
Endangered but largely overlooked mammal species, with potentially important implications for understanding its ecological requirements and thus informing future conservation management. However, additional analysis and some restructuring of the manuscript is required before it can be published.

Comment
The introduction is all a bit too specific, and makes the paper feel like it would only appeal to someone already interested in muntjacs. It would be much better to open the paper by framing the study in a much wider context, by starting off with an explanation about the importance of using data from the past (historical baselines, environmental archives) to provide new insights for conservation of threatened species, and the ways in which this can be / has been done -i.e. make the paper of interest to people beyond just muntjac specialists!

Response
We agree with this comment and the study would benefit from being placed in a wider research context. The introduction has been revised and expanded with the addition of the following section:

PAGE 1 lines 44-59
"Human activities continue to reduce mammal populations, geographic ranges and, ultimately, cause extinctions [1]. While the scale, rapidity and mechanisms driving recent anthropogenic impacts such as these are unprecedented [2-3], attempts to form effective conservation strategies are potentially hampered by a paucity of studies that consider biological communities from millennial as well as ecological (typically less than 50 years) timescales [2,[4][5][6][7]. Current studies and recent data (less than 100 years) characterise animal communities and their habitats only after, potentially, centuries or millennia of exploitation and modification by humans [7][8][9][10]. In this sense, a historical amnesia results in a shifting baseline syndrome where ecosystems, animal populations and their current geographic distribution are interpreted as 'natural' or pristine, where they are in fact degraded [2,9,11]. This issue is likely to be particularly problematic with rare, recently described and poorly-known mammals [7,11]. In this context, the potential of Quaternary archaeological and palaeontological data to provide longer timescale perspectives and benchmark evidence for biological conservation is increasingly being recognised and demonstrated [7,[9][10][11][12][13][14][15][16]. Here, we present just such a line of evidence and consider a poorly-known and Critically Endangered species of deer (Cervidae): the giant muntjac." The bibliography has been updated accordingly, references 1-16.
Revised manuscript RSOS-181461.R1 Stimpson et al. An 11,000-year-old giant muntjac subfossil from Northern Vietnam: implications for past and present populations 2

Comment Similarly, the previous suggestion that vuquangensis and gigas are the same species should be introduced in the introduction, rather than this important point being relegated to the discussionas your study provides further evidence of a past wider distribution of giant muntjacs, which supports the previous study.
Similarly, the idea that the name "gigas" can be "ignored" for now, even though it represents a senior synonym of vuquangensis, because "this revision has not yet been formally adopted", is a bit of a red herringthe way it can be adopted is by studies like this deciding to use the name "gigas" (it doesn't require a formal IUCN edict)as there is no worldwide arbiter of taxonomy. So, if the authors consider that vuquangensis and gigas are the same taxon, they should use the name gigas in this manuscript; and if not, then they should state why they think they're different.

Response
In light of the reviewer's comments, it is clear that the taxonomic discussion of Muntiacus vuquangensis/gigas requires clarification from the outset. We agree that, following the study of Turvey et al. (2016) . We therefore adopt the use of M. gigas, which we consider to be synonymous with M. vuquangensis.

Response
We agree and we sort available comparative measurements for M. gigas before submission of the manuscript and made renewed searches and appeals to colleagues following the reviewer's comments. We were successful in sourcing limited data for "M. vuquangensis" which as per the discussion above, is regarded as the same species. The original description of M. gigas fragmentary mandibles and maxilla are alluded to but no descriptions or measurements are presented (Wei et al. 1990). Subsequent records also appear to have focused on the identification and description of pedicles and antlers. To our knowledge there are no available published dental data with a confirmed ID for M. gigas: we are therefore grateful to R1 for supplying m2 and m3 length data for this taxon.
These data were incorporated as follows:

PAGE 7 lines 10-12
Comparison of m2 length (SLML) and m3 length (TLML) from the M. gigas specimen indicated no significant difference between these measures and the normative sample of M. muntjak. See also response to comment below:

Comment The lack of any statistical analysis to compare the new specimen with other muntjac samples (muntiacus, vuquangensis, gigas) is a major omission, and really has to be doneeven if just via something fairly straightforward such as t-tests or ANOVAs.
A statistical assessment of measurement differences, rather than just "eyeballing" the data, is essential.

Response
We agree that the study would benefit from statistical testing of observations of metric data. Given that the archaeological specimen is a single sample simple t-tests or ANOVA's would not be appropriate, given n = 1, we have used a normative comparisons approach (described below) of a single case (dental metrics from the mandible) to a normative sample (metric data sets from comparative taxa).

PAGE 3 lines 55-60 to PAGE 4 lines 3-11
The methodology has been updated accordingly: Statistical tests were performed to compare dental metrics of comparative museum specimens and published sources [41][42][43]. Comparison of dental metrics from the mandible, and in cases where sample sizes of comparative data were small, testing followed the procedures for normative comparisons [44][45][46][47], where metrics from a single case (e.g. HBC-27587) are compared with a normative sample generated from comparative data to test for differences from the estimated parameters of a given taxon (e.g. M. muntjak). Multivariate normative comparisons (m2 and m3, lengths and widths or a subset of these data depending on the availability of comparative data) were performed using a modified Hotelling's T 2 test, following Huizenga et al. [46]. Univariate normative comparisons were also performed with a modified t-test [44-45] with a step-down correction to control familywise false-positive error rate in multiple comparisons [47]. These approaches assume univariate/multivariate normality in the normative samples. Sample data sets were examined for departures from normality using the functions in PAST 3.20. Normative comparisons were performed using the E-clip, Multivariate  The comparative section (3.2 Morphology and metrics -comparisons) has been revised in the following areas: PAGE 5 lines 47-50 (comparison with C. pygargussee also response to comments from R2 "Capreolus", below) The m2 and m3 lengths of the archaeological specimen exceed those of small samples of Holocene specimens from Primoye [43] but these differences are not significant (SLML: n = 4; t = 1.117; P = 0.25, one-sided; TLML: n = 5; t = 1.291; P = 0.09, one-sided).

PAGE 6 lines 54-60 and PAGE 7 lines 3-25
The limited available measurements from larger-bodied Muntiacus spp., M. crinifrons (n = 2) and M. feae (n = 2) fall within the upper range of M. muntjak. Multivariate normative comparisons of m2 and m3 lengths and widths (SLML, SLMW, TLML, TLMW) of available data from these species (n = 2, in both cases) with sample data from specimens of M. muntjak from mainland Southeast Asia (n = 23) indicate no significant difference (table 3).
Available data from two specimens of "M.  (table 3). This test indicated that the metrics from Schaller's female specimen are marginally, but significantly, larger than the normative sample (table 3). Comparison of m2 length (SLML) and m3 length (TLML) from the M. gigas specimen indicated no significant difference between these measures and the normative sample of M. muntjak.
The dimensions of the m2 and m3 of HBC-27587 exceed all available comparative data from species within the tribe (figure 5). Comparisons of m2 and m3 lengths and widths (n = 23) and m3 lengths and widths (n = 32) of HBC-27587 indicate that these variables are significantly larger in the archaeological specimen, than in the sample data for M. muntjak (table 3). The body weight estimates derived from measurements of the mandible are also consistent with the trend in dental metrics and suggest a larger animal than the majority of Muntiacus spp. A mean estimate of approximately 40 kg for the specimen exceeds reported weights ranges of all taxa within the tribe bar one: M. gigas ( figure 6). While the dental dimensions of the Hang Boi specimen exceed available comparative data, given the range of variation within dental measurements in the genus (figure 5) there is no reason to suggest that the archaeological specimen would be an excessively or anomalously large specimen for this taxon. As such, there are no compelling grounds to posit a novel species and we refer specimen HBC-27587 to M. gigas (syn. vuquangensis). (table 3): One-sided (larger) 0.266 0.394

Minor comments:
-"sub-fossil" should be changed throughout the manuscript to "subfossil" (the more normal way of spelling this word) Changed to "subfossil" throughout the manuscript -"critically endangered" should be capitalised to "Critically Endangered", as this is a formal IUCN threat category. It also shouldn't be written in inverted commas (line 58) Changed to Critically Endangered, throughout.
-Line 112: "11, 778"spacing after the comma needs correcting Corrected - This section is revised and expanded:

PAGE 8 lines 36-43
The species has been recorded up to 1200 m above sea level (asl) but available data suggests the species is most likely to occur below 1000 m asl. Reliable estimates of lower elevational limits, however, are confounded by recent trends in habitat removal and over-hunting [31]. The Tràng An karst rarely approaches 200 m asl and the coastal plain that surrounds the massif, and dolines within it, are only a few meters above sea level. The specimen from Hang Boi indicates that giant muntjac occurred in a lowland karst setting and, as such, the current altitudinal range of the giant muntjac in the Annamites may not represent an ecological preference for habitats at higher altitudes, but rather a response to human activities.  (NHMUK 83.324) suggest similar dimensions to HBC-27587 (figure 5). Lengths and widths of the third molar are smaller than the archaeological specimen and fall into the upper size range of equivalent data for M. muntjak ( figure 5). The m2 and m3 lengths of the archaeological specimen exceed those of small samples of Holocene specimens from Primoye [43] but these differences are not significant (SLML: n = 4; t = 1.117; P = 0.25, one-sided; TLML: n = 5; t = 1.291; P = 0.09, one-sided).

REVIEWER 2
Morphologically, the concavity of the posterior margin of the mandibular ramus appears shallower but much wider (i.e extends much further dorsally to the condyle) in Capreolus than is suggested in the archaeological specimen. C. pygargus lacks anterior cingulids, as is the case with the archaeological specimen, but protoconids and hypoconids are much more angular and there is marked overlap of the metaconids and entoconids. Ectostylids are present on the lower molars, though are weakly developed, particularly on the m3, in comparison to HBC-27587 ( figure 3).
The genus Capreolus is associated with temperature climates at higher latitudes both in terms of extant populations and the wider fossil record in Eurasia [60]. In Pleistocene records of East Asia, C. pygargus does not appear as a component of the Ailuropoda-Stegodon fauna [61] and Holocene records (archaeological and historical, n = 51) are restricted to central and northern provinces in China, with no records of the species further south than Hubei [11]. As such, C. pygargus can also be reasonably discounted on biostratigraphic and as well as ecological grounds (see section 4.2).
Limited available data from Capreolus pygargus is also included in figure 5 and occlusal morphology of the m2 and m3 in now shown in figure 3.

Elaphodus: PAGE 6 lines 12-22
The tufted deer, Elaphodus cephalophus, is currently restricted to southern China (with historical records from eastern Myanmar). These deer are associated with montane forest habitats and apparently do not range into sub-tropical environments [ figure 6) and metrics of the m2 and m3 (figure 5) indicate that extant Elaphodus are relatively small in comparison to equivalent data from the archaeological specimen. Comparison of m3 lengths and widths (TLML, TLMW) with a sample of extant E. cephalophus (n = 9; this study, [42]) indicate that these dimensions are significantly larger in the archaeological specimen (T 2 = 26.498; P < 0.001; one-sided).

PAGE 7 lines 50-51
"This likely to be a complex issue to address at present and a matter of interpretation." corrected to "This is likely to be a complex issue to address at present and a matter of interpretation." Comment "find this sentence hard to understand, perhaps separating into two sentences would help." This section has been revised and clarified. Please also note that on review, the original "M. munjak" data set, erroneously contained data from M. feae and M. crinifrons. While it is unlikely that these small samples had a significant impact on the characteristics of the distribution of measurements (as discussed in the manuscript in the morphometric description of these species), it was not appropriate to include them and the Kruskal-Wallace tests were re-run. The changes to the outcome of these tests were marginal and subsequent interpretation was not affected. Furthermore, the data point for m3 length for M. gigas (kindly supplied by R1) was grouped with m3 lengths of M. vuquangensis (synonymous) and HBC-27587 (the archaeological mandible) as a sample for M. gigas, to compare with the Middle and Late Pleistocene data sets.

PAGE 8 lines 9-19
A comparison of Middle and Late Pleistocene m3 lengths from specimens attributed to "M. muntjak" with data from extant M. muntjak (n = 32; this study, Comment "I suggest to put in parentheses for the sake of consistency" PAGE 9 lines 29-30 "and civet, Parodoxurus hermaphroditus." changed to "and civet (Parodoxurus hermaphroditus)."

COMMENTS ON FIGURE 2
"is that meters above sea level?" "this figure shows less layers than given in Figure S1. Specification is needed." Figure 2 shows the upper 2 m of the stratigraphic section and heights relative to an arbitrary site datum (set at 500 m) rather than sea level. This issues are clarified in revised figure caption: The caption for table S1 is also amended for clarity. Table S1. AMS radiocarbon dates from the total excavated extent of the midden sequence from Hang Boi (HBC), Tràng An World Heritage Area, Northern Vietnam.

REVIEWER 3
The manuscript is well written and if the topic falls into the scope of the journal, it definitely is worth to be published.

Comment
Unfortunately I have not found any access to the supplementary tables and figures. Therefore, I could not comment some of the evidence used in the ms.

Response
We regret that the R3 could not gain access to the supplementary information, as we feel the contents address several of the reviewer's comments, below:

Comment
Substantial part of the arguments is based on the body mass estimation. The method is recalled from Janis (1990). Since the reference deals with a chapter from a book published almost 30 years ago, it may be difficult for many readers to get and read the methodology there. This may have two implications to the recent manuscript. First, the authors of the present study should describe in brief the principles of the method for those readers who do not have the cited book at hand.

Response
In light of the reviewer's methodological concerns that potential readers may not have access to the source material (Janis, 1990) and for the sake of transparency, the parameters of the regression equations and output (which were originally included in the in Supplementary Materials as table S2) have been included as table 2 in the revised manuscript. The methodology section is also revised: PAGE 3 lines 43-47 Body mass estimates were derived from six measurements of the specimen, following the least squares regression equations and percentage standard errors of estimate (%SEE) for the Cervidae of Janis [39; table 2].

Comment
Second, from a more recent literature (e. g., Mendoza, M., Janis, C. M. & Palmqvist, P., 2006. Estimating the body mass of extinct ungulates: a study on the use of multiple regression. J. Zool. 270, 90-101;De Esteban-Trivigno, S. & Köhler, M., 2011. New equations for body mass estimation in bovids: Testing some procedures when constructing regression functions. Mamm. Biol. 76,[755][756][757][758][759][760][761] it is apparent that some improvements of the method are available since 1990. Even if it was not applicable to the paleo-material available, it seems the body mass estimation based on multiple regression should be considered more carefully.

Response
While the methods, above, present refined means of estimating body mass, they are either practically (deal with characters or character sets not available to this study) or taxonomically (Bovidae, not Cervidae) inappropriate. To our knowledge, the equations of Janis (1990) remain the most comprehensive and robust treatment of body mass estimations for Cervidae (see also Suraprasit et al., 2016).

Response
A figure showing these measurements was (and is) included in the Supplementary  Information as figure S1.

Comment
Ad Figure 3 -Chinese water deer announced in the text (lines 196-198) is absent in the figure.

Response
Hydropotes now shown in figure 3 (see also response to R2 comments, above)
Correct. As discussed in section 2.2 of the manuscript (PAGE 3 lines 17-18): "The mandible was recovered in the western half of a 1 m x 1 m grid square (226/109) in a 10cm unit of excavation (5010) through context (5105)". Table S1 gives the excavation unit 5010 (within which the charcoal sample was recovered) rather than the context (5105): this is corrected to 5105 in the table, for clarity.
Comment. "Why not using here the same abbreviations as for Table S2 and Figure S1? Otherwise you have to explain that L and M are equal with SLML, SLMW, TLML, and TLMW"