Excellent manuscript in isotopic applications in archaeology because clearly, the authors have considered the assumptions related to the analytical techniques applied, animal physiology, and all the caveats and careful considerations that are necessary for properly using SI in archaeology. The authors successfully extracted as much information as they could from ungulate tooth enamel using carbon and oxygen isotopes adding more valuable environmental and ecological information in northern and northeastern Spain, regions where such records are growing, providing detailed scenarios for the region during the MP-UP transition. This manuscript is well-written and well-structured. Data are presented well in tables and figures.  Besides a few points that need clarification, the article should be accepted for publication. 

Line 95: the word “approach” is too general, be more specific.

Line 98: Refrain from using word correlate unless speaking about statistical correlations.

Fig. 2- why is only one circle around 40,000 years ago green and all the others black, needs to be explained in the caption.

This is a remarkable manuscript that deserves publication after major changes.

From my point of view there are some overinterpretations on the one hand, and some oversimplifications on the other hand. An important problem of paleoenvironmental reconstructions based only on archaeological remains, like this manuscript, is that the obtained data are not contextualized in an objective paleoclimatic and paleoenvironmental frame, in this case (northern) Iberia. This study based the paleoenvironmental reconstructions mainly on the archaeological data instead of comparing them with regional continuous paleoclimatic and paleoenvironmental records. I mean, authors suggested that this type of paleoenvironmental studies is key to understand past climate and human interactions. See for example abstract lines 18-19 or the introduction. Authors must keep in mind that the paleoenvironmental reconstruction that they have performed in these archaeological sites are “discontinuous points” in the paleoclimatic record of the Iberian Peninsula (see for example the chronologies in Fig.2). I mean, the most accurate climatic records for the studied periods are marine, lacustrine and speleothem records, where one can observe the “objective” fluctuations of past climate. Authors should acknowledge this point in the manuscript as well as compare/discuss with these records (there are many for the IP, for example Martrat et al., 2004; 2007; Pérez Mejías et al., 2019; Moreno et al., 2012; González-Sampériz et al., 2020; Camuera et al., 2019; 2022, among others). Some of them also discuss about vegetation changes in NE Iberia, close to the study areas, which would help authors contextualize and discuss their interpretations about the animal diets. Similarly, the speleothem records from the north of Iberia would help constrain temperature/precipitation patterns; there are many of these records in the Vasco-Cantabrian area. Other records from the Iberian Peninsula, the Mediterranean coast or the Iberian margin would show the general climatic patters for this period. Authors should discuss their data (agreement or disagreement) according to these continuous records in order to have a big picture of the paleoclimate and paleoenvironments in the studied period. Otherwise, authors are only comparing the archaeological sites from their two study areas and the work would only be of local interest. In fig. 2 authors included the d18O record of Greenland, but they did not discuss their data according to this record. As I mentioned before, they should compare and discuss the Iberian records instead, since there can be some temporal offsets between some of the events of the NGRIP curve and the Iberian/mediterranean records. Taking all this into account, objective 3 (line 96) is not totally achieved. In any case, my main concerns about this work come from the methodological approach.

Chronology: This a very important part of the study and should be presented in a subsection in section 2 or in the methodology (section 3). Please, explain deeply the absolute chronology of the levels where the ungulated remains were collected. Please, specify the dating method: ESR, OSL or 14C in Fig 2. Regarding 14C samples: it is not clear what radiocarbon curve was used to calibrate the 14C samples. Please, re-calibrate all the radiocarbon data with the latest 14C curve from 2020; there are important age shifts after the Holocene comparing the intCal2020 curve and previous calibration curves. Takin into account the confidence intervals of the ages (I suppose 2 sigma for 14C)->Line 98-99: “The chronological resolution in the study areas for this period allows us to correlate regional paleoenvironmental changes with global records”: this would be only true for the two sites younger than 30 ka, since the dates of the other sites might overlap stadial and interstadials (and their probability distributions are very large). What are the grey bands in Fig. 2, stadials? Please explain. In any case the chronology of the samples, according to Fig. 2, would only allow to differentiate between stadials/interstadials in the two youngest samples. What is the meaning of the green colour in one date at around 40 ka in Fig. 2 (Canyars)? What is the meaning of the dates (dots and bars) in Fig 2: Do they represent a single dating event, or a sum of distribution of various dates?, please explain this in the caption and in the main text. If it is the sum of distribution, did you use any statistic approach to obtain it (such as the ones that can be obtained from OxCAl software?) Do they represent the ages of the whole archeological sites or of the levels where the ungulated teeth were taken? This is very important to specify in the main text since the age of the remains could vary. Therefore, the caption of Fig. 2 should explain these details and there might be a (sub)section in the main text explaining the chronology of the levels where the remains were taken. This is crucial to validate the discussion of the manuscript

Methodology.

1. Authors mentioned that they did not carry out any pre-treatment to remove secondary carbonates, but did authors check the potential presence of secondary carbonates or the preservation of carbonates?. This is very important since all the results are based on these values (there is no data of d18O in phosphates). The physical cleaning that was carried out would not remove all the potential secondary carbonates. Secondary carbonates are very common in archeological contexts such as karstic caves (like the ones studied here), and would modify the isotopic composition of carbonates if they are not eliminated. This must be double checked before stating the sample preparation. Authors did not mention methods to double checked that the isotopic signal was the pristine one. They only mentioned in line 503-504: “The carbonate content in our samples, ranging from 3.9% to 8.9%, is similar to the proportion found in modern tooth enamel, suggesting no immediate indication of diagenetic alteration”. However, there is no explanation about the methodology used to calculate this percentage of carbonate. Authors should explain this in the methodology section and add the % of carbonate in a table (e.g. Table 2).

2. Regarding the potential treatment to remove the organic matter, authors said: Lines 145-151: “For this reason, in this work, most of the samples were not pretreated, except for the equid samples from Labeko Koba and Aitzbitarte III, and the cervids and equids from El Otero that were sampled and pretreated in the context of the initial project. Pretreatment followed was established by Balasse et al. (2002), where around 7 mg of powdered enamel was prepared and pretreated with 3% of sodium hypochlorite (NaOCl) at room temperature for 24 h (0.1 ml/mg sample), and thoroughly rinsed with deionised water, before a reaction with 0.1M acetic acid for 4 h (0.1 ml/mg sample) (equivalent protocol in Jones et al., 2019).” And afterwards in lines 453-456: “In the case of equid samples from the Vasco-Cantabrian region, it should be considered that they have been pretreated with a combination of NaClO and acetic acid, which could potentially affect the isotopic values. Samples after organic removal pretreatment can potentially show either higher or lower δ13C values and higher δ18O values based on previous experiments (Pellegrini and Snoeck, 2016; Snoeck and Pellegrini, 2015)”. So, my doubt is: why did authors treat whole batches of samples to evaluate these “side effects” instead of applying both protocols (with pretreatment and without pretreatment) to aliquots of the same samples? Although they finally ended up that “the influence of the pre-treatment on our samples is deemed to be limited.“, this was a risk, and now they cannot be 100% sure about this potential influence. Was there any reason to measure the samples where the organic matter was removed in a different IRMS. If yes, please, explain.

3. Line 182-196 and throughout the calculations and the manuscript: Authors referred to E* as the fractionation factor. The symbol E is traditionally the enrichment factor, not the fractionation factor (alpha). E= (alpha-1)x1000‰. So, what factor have authors applied eventually: fractionation or enrichment factors? Are these factors mixed in the text and in the calculations? This has to be clear, and if the factors are wrongly applied (enrichment instead of fractionation factor), correct the calculations. Although both factor are related (E= (alpha-1)x1000‰.) the obtained results would differ, and thus, the derived potential interpretations.

4. Lecuyer et al. (2021) performed the calculations to correct the effect of atmospheric CO2 (difference of 1‰ and CO2 concentration) for the LGM; so, these specific CO2 corrections can only be applied for the LGM, but in the present manuscript there are no samples for the LGM (23-19 ka). In addition, the correction for the isotopic composition of atmospheric CO2 should be done specifically for each age of the studied samples, instead of using a general average -7‰: as authors mentioned, a variation of a ca. 1‰ would imply a change in 150mm of precipitation. Check for example the isotopic composition of atmospheric CO2 reconstructed from ice records for the Late Pleistocene that ranges from -7 to -6.5 ‰ (Eggleston et al., 2016: Paleoceanography, among others). So, please, apply age-specific CO2 corrections. I mean, when you are quantifying climatic variables, you should reduce the potential error sources. These errors increase by applying a general unspecific correction for all the data (the same affirmation could be applied to all corrections of the isotopic data in the manuscript).

5. Authors proposed the above-mentioned (oversimplified) correction for the change in the isotopic composition atmospheric CO2, but they did not apply any correction related to the change of the isotopic composition of the sea-water during the Late Pleistocene, which is the main moisture source for rainfall (I would not mention the moisture and precipitation due to inland evaporation/recycling in the Iberian Peninsula, even during cold periods (Krklec and Domínguez-Villar, 2014), in order to simplify this interpretation). The global isotopic composition of the rain during colder/warmer periods (glaciar/interglaciar, stadial/interstadial) differs, not only due to the isotopic fractionation caused by temperatures, but also due to the accumulation/release of the lighter water isotopes in the ice sheets/glaciers during cold/warmer periods (Dansgaard, 1964), among others factors affecting the global isotopic composition of sea waters. Therefore, in order to obtain a reliable isotopic data related to precipitation, the obtained dO18 values has to be corrected to remove this effect. See for example, Niedermeyer et al. (2010) or Garcia-Alix et al. (2021) approach to correct past hydrogen isotopes from vegetation, or even Fernández-García et al (2020) for fossil mammals in the studied period of this paper.

6. Where did these obligate drinkers drink (water source)? Directly from the rain? Ponds? Lakes? Unless they directly drink precipitation waters (oversimplification), this would imply more isotopic fractionation and would also mask the temperature signal. This is even more important in the studied glacial period, and especially in the stadials? Apart from the potential enhanced rain evaporation due to low atmospheric moisture in glacial times (Dansgaard 1964), an increasing evaporation in lakes, ponds (and even in vegetation->enhanced evapotranspiration) during dry periods have been demonstrated by different isotopic studies in carbonates from freshwater gastropods, bivalves, or ostracods and even from leaf wax isotopes of freshwater and terrestrial plants. Please clarify this issue, and explain this constrain in the methodology since it would affect the reconstructed temperatures.

7. Precipitation source (North Atlantic Oscillation modes and Mediterranean dynamics). In the Iberian Peninsula, the isotopic composition of precipitation is highly affected by the moisture source in the present - and in the past - (Araguás y Díaz Teijeiro, 2005, Celle-Jeanton et al., 2001; Domínguez-Villar et al., 2013; Krklec and Domínguez-Villar, 2014; Moreno et al., 2010; 2012; 2014; 2021, Toney et al., 2020; García-Alix et al., 2021; Schirrmacher et al., 2020, among others). Therefore, temperature-isotope fractionation equations would not work that well to reconstruct past temperatures. When proper analysis of the isotopic signal of precipitation are performed in N Iberia, there are sampling stations (precipitation, temperatures, isotopes, and moisture sources) where the isotopes from precipitation do not correlate well with temperatures (Moreno et al., 2021). This thoroughly study of the isotopic composition of precipitation in northern Spain ended up with “although important, air temperature only explains part of the observedδ¹⁸O_pvariability and is therefore not the only control.” This issue is even more important in coastal areas, as the ones studied in this paper, and even more in the Mediterranean coast (at present there is some influence of amount effect in the Mediterranean areas of Catalonia; Moreno et al., 2021). Therefore, in the best oversimplified case-scenario (not considering previous comments 5 and 6, and admitting large errors due to these potential source and amount effects) if we would want to calculate temperatures from the isotopic values of precipitation we would need an equation for the Vasco-Cantabrian region, and another one for NE-Mediterranean Iberia (different precipitation pattern and forcing). And even more, since atmospheric patterns, and therefore, moisture sources, are not the same during the warm and cold seasons (see the above-mentioned studies), specific equations for cold and warm seasons should be applied to reconstruct “summer” and “winter” temperatures.

8. “MAT was calculated from the d18O mean value between summer and winter in each tooth before modeling to reduce associated error”; However in caption Table 4: “For some profiles with an unclear seasonal shape, MATs were deduced from the original average of teeth without a seasonal profile”. So, what is the correct methodology?, In any case, according to the methodology section, MAT was calculated before modeling to reduce associated error, but summer and winter temperatures after the inverse modelling?. This reasoning is not clear to me: Is there no associated error in the transformation for summer and winter temperatures? This is why there are some odd values, for example, sample AXL60 MAT 12ºC, ST 20.4ºC and WT 10.8ºC (only 1.2ºC colder than the MAT).

9. The reconstructed meteoric waters are different depending on the species, even in the same level, and therefore, reconstructed temperatures also differ. I’m aware of the different ecological behaviors of the different species, but the MAT should be, at least close. There are also some discrepancies in specimens of the same species in the same archaeological levels.

10. The general comparison between the isotopic composition of the faunas of the archaeological sites of both areas is not objective since in the Vasco-Cantabrian area there are remains from 80 ka to 18 ka approx. (14 sections from 5 archaeological sites), but in the Catalonian area there is only one site at around 40 ka (it could have been coeval to the HS4, a period especially cold and dry). Thus, the general comparisons that appeared in some parts of the manuscript between two areas (not the ones specific at ca. 40 ka) are not balanced.

11. What is the error associated with the different equations Eq. 1-9? Authors have to take into account the different errors (not only the standard deviations) that are being accumulated in each equation and also plot and mention them in the tables and in the text. For example, a MAT of 15ºC +/- 0.5 ºC would be accurate, but if the error was +/- 4ºC the interpretation would be more open. This is even more important in this work since the oxygen isotopes were measured in carbonates, not in phosphates, and two more equations were used to converter the d18O data of the carbonates to d18O of phosphates. Authors said: lines 277-278: “In these estimations, the associated error from converting δ18Ophos to MAT is enlarged by the uncertainty derived from the transformation of δ18Ocarb (VPDB) to δ18Ophos (VSMOW)”. Therefore, all the errors should be calculated (for both O and C), included in the plots, tables and when describing the results.

12. Inverse Modelling. I think that the correct reference for the inverse modeling (main text and appendix D) is Passey et al. (2005) but this one “Passey, B.H., Cerling, T.E., Schuster, G.T., Robinson, T.F., Roeder, B.L., Krueger, S.K., 2005. Inverse methods for estimating primary input signals from time-averaged isotope profiles. Geochim. Cosmochim. Acta 69, 4101–4116.” Reviewing this paper, I noted that the reconstructed profiles showed mostly the same trends/changes as the original isotopic data. However, in the case of the reviewed manuscript sometimes these reconstructed profiles exhibited opposite patterns/trends from the original isotopic values. I’m not familiar with these kinds of transformations, but: is this common? This is an extra transformation for the data that would be used to calculate absolute climatic parameters, and thus, and extra potential error source. Could authors double check these calculations?. In addition, in the original paper from Pasey et al. (2005), this inverse modeling was also applied to the carbon isotopes. Why authors did not apply this correction also to the carbon isotopes of the sequential sampling?

Minor comments:

The first paragraph in the introduction and the first lines in the abstract deal with the importance of these kinds of studies to understand the human evolution in this region, but eventually, this is not discussed in the manuscript according to the obtained data.

First sentences of the introduction (lines 38-43): please add references. There are interesting papers dealing with this issue in the Iberian Peninsula and in Europe: Neanderthal-AMH-climate change.

Lines 49-50. It is ok, but authors are not including some information (and significant climate-related references) in the area dealing with continuous paleoclimatic records, and they only focused on the data obtained from archeological sites (whose paleoenvironmental record is not that continuous); so, line 75 is not summarizing the multiproxy studies in this area.

Lines 80-87: add references

Regarding the fossil sites that authors call “mediterranean”. Since the rest of the fossil sites are in “northeastern Iberia”, the term “Mediterranean area” is very open and do not specifically identify the studied site, I would say NE Iberia?

There are some issues with the chronology/dates. For example, line 21 abstract: 80 to 15,000 cal BP. Do you mean 80 ka or ky, right? Taking into account the study period, and the accuracy of the dates, I would not use “yr”, I would use ky or ka. In addition, authors should round the dates in the text to the nearest hundreds (eg, line 601: 41,136 to 38,570 cal yr BP: 41.1 ka to 38.6 ka). This accuracy does not make sense in the studied period due to the uncertainty of the measurements.

Results: please, add the ages (in ka and the different technocomplexes) to the subsection headings of the different archaeological sites, otherwise one has to check Fig. 2 for each site.

I understand the structure of the result description, but the mixed description of the isotopes from different levels of the same archaeological sites, which sometimes have 10 ka of difference between them, is rare to me.

Line 325: MATAs=1-8/-2.1ºC? do you mean 1.8?

Eq 10: P value <?

All figures with the data plotted according to the chronology. El Castillo 21A appears after Axlor III; but according to the chronology this site is previous (Fig. 2). The same would happen between some levels of Labeko Kova and Canyars-I. Is this correct? In this case, arrange the sites with the real chronological order even though they belong to different archaeological sections.

When speaking about range of values (temperatures, isotopes, precipitation), sometimes the lowest values are mentioned before and other times the other way around. Be consistent and cite the lowest values first (Eg, line 404  MATAs).

Table 3: I suppose that the different group of data (rows) are related to the three groups of specimens. Right? Please, specify the taxon groups in the tables.

I do not understand this sentence. Line 510-511: “Based on these arguments, it is suggested that the non-sinusoidal δ18O signal observed in some individuals is likely attributed to the preservation of the original isotopic signature from water input.“ This sentence would suggest a bad preservation of the original water composition when a sinusoidal pattern is present?. However, authors explained afterwards some reasons related to the ecology of the specimens. Please, clarify this.

Section 5.4. Regarding the d18O of meteoric waters. Authors compare (as a whole) their reconstructed d18Omw with the current values in the area and they ended up with similar ranges. Keep in mind that the reconstructed values correspond to the last 80 ka under glacial conditions (different temperatures, precipitation amount, and in some cases, moisture source)

Sometimes the different d13C and d18O have subscripts with the meaning of the isotopic values (eg, δ13Ccarb, δ18Ocarb), but another times this is not indicated. This is confusing. Please, add always the subscripts explaining the meaning of the isotopic values.

I miss a figure summarizing the chronological evolution of reconstructed temperatures and precipitation (with the associated accumulated errors) and comparing them with other continuous records of precipitation or temperatures or vegetation from the Iberian Peninsula, Mediterranean coast or Iberian margin.

Title: ecological evolution of what? This is very general: I would say ecological evolution of ungulate fauna….

Please, use always the symbol delta instead of d, there are some “ds” throughout the manuscript.

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