Parallels, differences and lessons: a comparison of the management of foot-and-mouth disease and COVID-19 using UK 2001/2020 as points of reference

Foot-and-mouth disease (FMD) is an extremely infectious viral infection of cloven-hoofed animals which is highly challenging to control and can give rise to national animal health crises, especially if there is a lack of pre-existing immunity due to the emergence of new strains or following incursions into disease-free regions. The 2001 FMD epidemic in the UK was on a scale that initially overwhelmed the national veterinary services and was eventually controlled by livestock lockdown and slaughter on an unprecedented scale. In 2020, the rapid emergence of COVID-19 has led to a human pandemic unparalleled in living memory. The enormous logistics of multi-agency control efforts for COVID-19 are reminiscent of the 2001 FMD epidemic in the UK, as are the use of movement restrictions, not normally a feature of human disease control. The UK experience is internationally relevant as few countries have experienced national epidemic crises for both diseases. In this review, we reflect on the experiences and lessons learnt from UK and international responses to FMD and COVID-19 with respect to their management, including the challenge of preclinical viral transmission, threat awareness, early detection, different interpretations of scientific information, lockdown, biosecurity behaviour change, shortage of testing capacity and the choices for eradication versus living with infection. A major lesson is that the similarity of issues and critical resources needed to manage large-scale outbreaks demonstrates that there is benefit to a ‘One Health’ approach to preparedness, with potential for greater cooperation in planning and the consideration of shared critical resources.


Do you have any concerns about statistical analyses in this paper? If so, please specify them explicitly in your report. No
It is a condition of publication that authors make their supporting data, code and materials available -either as supplementary material or hosted in an external repository. Please rate, if applicable, the supporting data on the following criteria.

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Comments to the Author This is a very interesting manuscript which is very detailed with a huge depth and range of information presented. To have its greatest impact for a general reader this reviewer feels that it needs to be a much shorter and concise document. The following are some suggestions 1. Table 1 is very effective. A number of Tables with comparisons between FMD and COVID19 under a number of different heading would greatly help the reader to understand the range of problem areas in the practice of One Health. A list of current data sources would also be helpful.

2.
While the authors are very clear about the outcome of the inquiry into FMD2001, it is lost in the detail of the manuscript. The lessons learned, which can all be applied to COVID 19 should form a central theme of the manuscript and be included in the abstract. 3.
The similarities in the lack of test capacity must be highlighted. While COVID 19 is considered a new virus, it is related to other SARS virus so the lack of investment in the development of accurate test capacity was not learned from FMD but must be learned for the next animal or human pandemic. 4.
While the authors report the percentage of the total heard that were culled, some standardisation of the death rate such as the proportion of the population of England who died from COVID 19 rather than the actual numbers should be provided for a given time (April 30th 2020). Comparison of crude numbers, as currently presented in the media has little meaning.

5.
A short section on the priorities for Public Health or One Health Preparedness, based on lessons learned from both Epidemics would also be helpful. 6. Table 2 is cited in the conclusion but I feel this is Table 1.
Because the paper switches between a UK and global focus, it is unclear to me what the primary research question is. If the primary research question is "What can we learn by comparing the current pandemic to the 2001 FMD epidemic?" then the paper must be revised to encompass the broader management strategies and incentives of FMD and COVID and provide a more balanced comparison of the two diseases. I do not need to compare COVID19 to FMD to know that early detection, testing, and movement restrictions are needed prevent spread and limit infections. But that is not to say that there are interesting lessons to be learned from the two. While some of the important details need to be added to more fully describe the 2001 epidemic, there is a story about the initial seeding and effectiveness of management efforts in regions that were overwhelmed or not by the number of FMD cases. In addition, there are some interesting questions regarding the global management of FMD, and the trade offs associated with eradicating FMD in developing countries vs. welfare, that will be applicable to future COVID19 management.
If the primary research question is "Can we use the 2001 FMD epidemic to evaluate if the United Kingdom improvement its biosecurity and management policies for facing new epidemics?" then the paper is in better shape. That being said, much of the paper will need to be reworked and the authors will need to decide on when to focus on the UK versus the global story.
Regardless of the editorial decision, I believe that there need to be a number of changes to improve the manuscript. First, the scale of the study -particularly that of the 2001 epidemicneeds to be clarified. The authors flip back and forth between management within the United Kingdom and global management of FMD in 2001, which is confusing and changes what we can say about disease management. The World Organization of Animal Health maintains a set of five disease-free categorization ranging from disease-free, no vaccinations (low risk) to not diseasefree (high risk). Countries with and without FMD face very different decisions in how they manage disease risk within their own countries but also on the global market. Questions of welfare and tradeoffs within developed and developing countries are important for FMD management (see the 2012 Global FMD Control Strategy) and will be for COVID19.
In this same vein, the nature of disease-free designations, and how they impact the different management strategies is important. Understanding why people choose their management decisions and how they are incentivized or compensated (maybe too well for the 2001 FMD epidemic) matters for the current management of an epidemic but also for how they will view and react to future risks of disease.
The temporal scales of the epidemics are also important. Where do each of the "phases" in the manuscript fall on the epidemic curves for each? Could the authors include a figure of each epidemic labeling the phases? There are some inconsistencies in the timing between the phases, such as testing centers being overwhelmed (which is in phase 1 but could easily fall in phase 2 and 3, no?).
Finally, if we are going to compare the two diseases, then it should be done so in a complete fashion. Many of the reasons that FMD is such a pain to manage are mostly ignored. Specifically, FMD infects virtually every cloven hoofed animal in the world, can be spread by multiple pathways with different animals exhibiting different susceptibilities and pathways of viral shedding, the presence of carrier animals, and wild disease reservoirs. While I understand that this will cause the comparison to diverge, it is important to remain transparent and open with the science of the two diseases.
I am including a number of other comments and suggestions embedded in the attached .pdf file. Please let me know if they are not viewable.

27-May-2020
Dear Dr Sumption: Your manuscript has now been peer reviewed and the reviewers' comments (not including confidential comments to the Editor) are included at the end of this email for your reference. As you will see, the reviewers think the article has the potential to be a useful contribution but they have raised some concerns with your manuscript that would need addressing first. Perhaps the most important are to clarrify the scope (just UK or global?) and be more up-front about the (big) differences between the diseases --see referee 3's comments in particular. However, because the referees have been quite specific in their comments, and these look to me to be achievable, I would like to invite you to revise your manuscript to address them.
We do not allow multiple rounds of revision so I urge you to make every effort to fully address all of the comments at this stage. If deemed necessary, 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. Please note that we cannot guarantee eventual acceptance of your manuscript at this stage. If the lessons are only for the UK and the differences between the diseases make extrapolation from FMD to COVID19 more tenetative, then the case for publication in Proceedings B seems less compelling.
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Thank you for submitting your manuscript to Proceedings B; we look forward to receiving your revision. If you have any questions at all, please do not hesitate to get in touch.

Best wishes, Innes Cuthill
Prof. Innes Cuthill Reviews Editor, Proceedings B mailto: proceedingsb@royalsociety.org Reviewer(s)' Comments to Author: Referee: 1 Comments to the Author(s) This is a very interesting manuscript which is very detailed with a huge depth and range of information presented. To have its greatest impact for a general reader this reviewer feels that it needs to be a much shorter and concise document. The following are some suggestions 1. Table 1 is very effective. A number of Tables with comparisons between FMD and COVID19 under a number of different heading would greatly help the reader to understand the range of problem areas in the practice of One Health. A list of current data sources would also be helpful. 2. While the authors are very clear about the outcome of the inquiry into FMD2001, it is lost in the detail of the manuscript. The lessons learned, which can all be applied to COVID 19 should form a central theme of the manuscript and be included in the abstract. 3. The similarities in the lack of test capacity must be highlighted. While COVID 19 is considered a new virus, it is related to other SARS virus so the lack of investment in the development of accurate test capacity was not learned from FMD but must be learned for the next animal or human pandemic. 4. While the authors report the percentage of the total heard that were culled, some standardisation of the death rate such as the proportion of the population of England who died from COVID 19 rather than the actual numbers should be provided for a given time (April 30th 2020). Comparison of crude numbers, as currently presented in the media has little meaning. 5. A short section on the priorities for Public Health or One Health Preparedness, based on lessons learned from both Epidemics would also be helpful. 6. Table 2 is cited in the conclusion but I feel this is Table 1.
Referee: 2 Comments to the Author(s) I enjoyed reading the article, which I found interesting and informative. However, I have a few remarks that should be considered in a revised version as follows: 1) The title should probably include reference to the "the UK" 2) Please compare estimates of R0 for both diseases. Can you say anything about the severity of the diseases? 3) While COVID-19 spread rapidly in the UK following an exponential growth pattern, the 2001 FMD epidemics spread following a slower growth pattern in the number of premises infected (see https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6132414/) 4) Please display the epidemic curves for both epidemics including a timeline of the key interventions Referee: 3 Comments to the Author(s) The manuscript "Foot-and-mouth disease in 2001 and COVID19 in 2020 : Parallels and differences in epidemic management" attempts to compare the characteristics of the 2001 footand-mouth disease epidemic with the current pandemic. I admit that my initial gut reaction to the abstract was a bit off-putting (I was afraid of a paper about COVID for the sake of COVID), but I found the paper an interesting read. Indeed, there are some similarities between the two: the presence of an aerosol-based infection pathway, the importance of testing for detection, and restriction of local, regional, and international movements. However, there are gross differences that make the comparison less compelling: many of the disease characteristics are quite different (e.g. FMD affects multiple species, with different levels of susceptibility and transmission routes, the presence of carrier species and wild reservoirs), sources of cost in the disease (FMD rarely kills -the main cost is in the trade response it induces), and obvious differences in management (we cannot cull people). Many of these are left out of the paper.
Because the paper switches between a UK and global focus, it is unclear to me what the primary research question is. If the primary research question is "What can we learn by comparing the current pandemic to the 2001 FMD epidemic?" then the paper must be revised to encompass the broader management strategies and incentives of FMD and COVID and provide a more balanced comparison of the two diseases. I do not need to compare COVID19 to FMD to know that early detection, testing, and movement restrictions are needed prevent spread and limit infections. But that is not to say that there are interesting lessons to be learned from the two. While some of the important details need to be added to more fully describe the 2001 epidemic, there is a story about the initial seeding and effectiveness of management efforts in regions that were overwhelmed or not by the number of FMD cases. In addition, there are some interesting questions regarding the global management of FMD, and the trade offs associated with eradicating FMD in developing countries vs. welfare, that will be applicable to future COVID19 management.
If the primary research question is "Can we use the 2001 FMD epidemic to evaluate if the United Kingdom improvement its biosecurity and management policies for facing new epidemics?" then the paper is in better shape. That being said, much of the paper will need to be reworked and the authors will need to decide on when to focus on the UK versus the global story.
Regardless of the editorial decision, I believe that there need to be a number of changes to improve the manuscript. First, the scale of the study -particularly that of the 2001 epidemicneeds to be clarified. The authors flip back and forth between management within the United Kingdom and global management of FMD in 2001, which is confusing and changes what we can say about disease management. The World Organization of Animal Health maintains a set of five disease-free categorization ranging from disease-free, no vaccinations (low risk) to not diseasefree (high risk). Countries with and without FMD face very different decisions in how they manage disease risk within their own countries but also on the global market. Questions of welfare and tradeoffs within developed and developing countries are important for FMD management (see the 2012 Global FMD Control Strategy) and will be for COVID19.
In this same vein, the nature of disease-free designations, and how they impact the different management strategies is important. Understanding why people choose their management decisions and how they are incentivized or compensated (maybe too well for the 2001 FMD epidemic) matters for the current management of an epidemic but also for how they will view and react to future risks of disease.
The temporal scales of the epidemics are also important. Where do each of the "phases" in the manuscript fall on the epidemic curves for each? Could the authors include a figure of each epidemic labeling the phases? There are some inconsistencies in the timing between the phases, such as testing centers being overwhelmed (which is in phase 1 but could easily fall in phase 2 and 3, no?).
Finally, if we are going to compare the two diseases, then it should be done so in a complete fashion. Many of the reasons that FMD is such a pain to manage are mostly ignored. Specifically, FMD infects virtually every cloven hoofed animal in the world, can be spread by multiple pathways with different animals exhibiting different susceptibilities and pathways of viral shedding, the presence of carrier animals, and wild disease reservoirs. While I understand that this will cause the comparison to diverge, it is important to remain transparent and open with the science of the two diseases.

Quality of the paper: Is the overall quality of the paper suitable? Good
Is the length of the paper justified? Yes Should the paper be seen by a specialist statistical reviewer? No Do you have any concerns about statistical analyses in this paper? If so, please specify them explicitly in your report. No It is a condition of publication that authors make their supporting data, code and materials available -either as supplementary material or hosted in an external repository. Please rate, if applicable, the supporting data on the following criteria.

Do you have any ethical concerns with this paper? No
Comments to the Author (a) A population adjusted rate for the impact of SARS COV-2 rather than raw data for the number of deaths/infections/positive cases would greatly add to usefulness of the data for future pandemics. This is particularly important as testing has been so limited to date. Similarly a population based approach to the size of the cul for all animals is required.
(b) While the Data provided in Figure 2 is from other sources it would be helpful if the onset of each outbreak lined up at the start of the outbreaks

Review form: Reviewer 3
Recommendation Accept with minor revision (please list in comments)

Scientific importance: Is the manuscript an original and important contribution to its field? Good
General interest: Is the paper of sufficient general interest? Good Quality of the paper: Is the overall quality of the paper suitable? Excellent

Should the paper be seen by a specialist statistical reviewer? No
Do you have any concerns about statistical analyses in this paper? If so, please specify them explicitly in your report. No It is a condition of publication that authors make their supporting data, code and materials available -either as supplementary material or hosted in an external repository. Please rate, if applicable, the supporting data on the following criteria.

Do you have any ethical concerns with this paper? No
Comments to the Author Overall, I find the paper much improved. It does a much better job at presenting a clear, up-front and accurate comparison of FMD and Covid-19, and the strengths/limitations on comparing them. My remaining concerns are primarily editorial, with several clarifications and typos. I move for acceptance pending (very) minor revisions.
Lines 17-18. The wording "… have experience of national scale epidemic crises of both diseases" is clunky. Reword to, "…have experienced epidemic crises on the national scale." Lines 18-19. Reword to "… FMD and COVID-19, with respect to their management including…" Line 43. In this section, you do not include a comparison to SARS-CoV-2. Consider renaming the header.
Lines45-49. Please include the citations in the text. (They are already in the References section from Table 1, so including them here will not increase the word length of the paper.) Line 48. Typo. Correct to "… and by ingestion, pigs being more readily…" Line 49. Typo. Add such that "… to ruminants, which occurs…" Lines 50-51. "… significant farm-to-farm spread can continue after imposition of complete animal movement standstills." It is worth noting that this depends on the spatial layout of farms. The fragmented farm landscapes characteristic of Europe is very different from the single, large tracks spread out and/or isolated in the United States system. Line 59. Correct to "… mid-April 2001, and at the height …" Lines 51-52. Because you talk about the large costs of the disease after, it is worth briefly mentioning them here, e.g. the culling of over 2 million heads of livestock (Sobrino and Domingo, 2001) and total costs of about £6.5 billion (Thompson et al., 2002) (doi's below).
Harapan et al., 2020: 10.1016/j.jiph.2020.03.019 Lines 99-102. "One driver of wildlife consumption… which occurred immediately before COVID19 emergence." Consider deleting this sentence. We are fairly certain that Covid-19 came from bats, with an intermediary host between bats and people. This sentence does not contribute much and is, if anything, misleading of the complex process inherent to Covid-19.
Line 119. Change to, "… resources. Such models have been built…" Line 125. Since this is a quote, add the reference number to the end of the sentence such that the text reads, "… reappraise their strategies and operations" [26]".
Line 138. "Information on the movements of livestock and where they are kept are routinely collected in the EU, especially for movement between countries." Could you add a reference where the reader could access information related to this?
Lines 178-180. "… the EU member states cooperate… centrally agreed controls on movements of animals and animal products across borders." Could you direct the reader to what I assume is a WTO plan? Line 181. Is "delay" the right word? Could you clarify/be a bit more specific? I am fairly certain that you mean "delay" [spread], but my mind went immediately to "delay" [introduction], which was confusing.
Line 195. Typo. Include a space such that "… in 2001 was a rationale…" Line 219. Run-on sentence. Correct to, "… like COVID19. This is a challenge." Lines 234 and 237. What do you mean by the phrases "the same periods of 2001" and "the denominators"? I do not understand. Please rephrase.
Line 242. Run-on sentence. Correct to, "… automatic culls. However, by then the scale…" Line 264. It is worth pointing out the approximate percentage of the population that must be immune in order to achieve herd immunity (it should be >80%). This will greatly strengthen the argument of the "high burden of disease". Line 289. You can cite Shanafelt and Perrings (2017) for a map/table of the global distribution disease-free/not disease-free countries for FMD.
Line 339. Typo. Rephrase to "… national crisis to plan for common, critical reserves…" Line 341. Tweak to "… democracies, and where such approaches…" Line 342. Add such that the paper reads, "… there may be many parallel local, national, and international veterinary…" Lines 345-346. To be consistent, use quotations around "One Health".

05-Oct-2020
Dear Dr Sumption I am pleased to inform you that your manuscript RSPB-2020-0906.R1 entitled "Parallels, differences and lessons: a comparison of the management of foot-and-mouth disease and COVID19 using UK 2001/2020 as points of reference" has been accepted for publication in Proceedings B.
The referees are happy with the revisions and have recommended publication, but also suggest some further minor revisions. Therefore, I invite you to respond to the referees' comments and revise your manuscript. Because the schedule for publication is very tight, it is a condition of publication that you submit the revised version of your manuscript within 7 days. If you do not think you will be able to meet this date please let us know.
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Once again, thank you for submitting your manuscript to Proceedings B and I look forward to receiving your revision. If you have any questions at all, please do not hesitate to get in touch.

Best wishes, Innes
Prof. Innes Cuthill Reviews Editor, Proceedings B mailto: proceedingsb@royalsociety.org Referee: 1 Comments to the Author(s) (a) A population adjusted rate for the impact of SARS COV-2 rather than raw data for the number of deaths/infections/positive cases would greatly add to usefulness of the data for future pandemics. This is particularly important as testing has been so limited to date. Similarly a population based approach to the size of the cul for all animals is required.
(b) While the Data provided in Figure 2 is from other sources it would be helpful if the onset of each outbreak lined up at the start of the outbreaks Referee: 3 Comments to the Author(s) Overall, I find the paper much improved. It does a much better job at presenting a clear, up-front and accurate comparison of FMD and Covid-19, and the strengths/limitations on comparing them. My remaining concerns are primarily editorial, with several clarifications and typos. I move for acceptance pending (very) minor revisions.
Lines 17-18. The wording "… have experience of national scale epidemic crises of both diseases" is clunky. Reword to, "…have experienced epidemic crises on the national scale." Lines 18-19. Reword to "… FMD and COVID-19, with respect to their management including…" Line 43. In this section, you do not include a comparison to SARS-CoV-2. Consider renaming the header.
Lines45-49. Please include the citations in the text. (They are already in the References section from Table 1, so including them here will not increase the word length of the paper.) Line 48. Typo. Correct to "… and by ingestion, pigs being more readily…" Line 49. Typo. Add such that "… to ruminants, which occurs…" Lines 50-51. "… significant farm-to-farm spread can continue after imposition of complete animal movement standstills." It is worth noting that this depends on the spatial layout of farms. The fragmented farm landscapes characteristic of Europe is very different from the single, large tracks spread out and/or isolated in the United States system. Line 59. Correct to "… mid-April 2001, and at the height …" Lines 51-52. Because you talk about the large costs of the disease after, it is worth briefly mentioning them here, e.g. the culling of over 2 million heads of livestock (Sobrino and Domingo, 2001) and total costs of about £6.5 billion (Thompson et al., 2002) (doi's below).
Harapan et al., 2020: 10.1016/j.jiph.2020.03.019 Lines 99-102. "One driver of wildlife consumption… which occurred immediately before COVID19 emergence." Consider deleting this sentence. We are fairly certain that Covid-19 came from bats, with an intermediary host between bats and people. This sentence does not contribute much and is, if anything, misleading of the complex process inherent to Covid-19.
Line 119. Change to, "… resources. Such models have been built…" Line 125. Since this is a quote, add the reference number to the end of the sentence such that the text reads, "… reappraise their strategies and operations" [26]".
Line 138. "Information on the movements of livestock and where they are kept are routinely collected in the EU, especially for movement between countries." Could you add a reference where the reader could access information related to this?
Lines 178-180. "… the EU member states cooperate… centrally agreed controls on movements of animals and animal products across borders." Could you direct the reader to what I assume is a WTO plan? Line 181. Is "delay" the right word? Could you clarify/be a bit more specific? I am fairly certain that you mean "delay" [spread], but my mind went immediately to "delay" [introduction], which was confusing.
Line 195. Typo. Include a space such that "… in 2001 was a rationale…" Line 219. Run-on sentence. Correct to, "… like COVID19. This is a challenge." Lines 234 and 237. What do you mean by the phrases "the same periods of 2001" and "the denominators"? I do not understand. Please rephrase.
Line 242. Run-on sentence. Correct to, "… automatic culls. However, by then the scale…" Line 264. It is worth pointing out the approximate percentage of the population that must be immune in order to achieve herd immunity (it should be &gt;80%). This will greatly strengthen the argument of the "high burden of disease".
Line 289. You can cite Shanafelt and Perrings (2017) for a map/table of the global distribution disease-free/not disease-free countries for FMD.
Line 339. Typo. Rephrase to "… national crisis to plan for common, critical reserves…" Line 341. Tweak to "… democracies, and where such approaches…" Line 342. Add such that the paper reads, "… there may be many parallel local, national, and international veterinary…" Lines 345-346. To be consistent, use quotations around "One Health".

12-Oct-2020
Dear Dr Sumption I am pleased to inform you that your manuscript entitled "Parallels, differences and lessons: a comparison of the management of foot-and-mouth disease and COVID19 using UK 2001/2020 as points of reference" has been accepted for publication in Proceedings B.
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Response to referees
We appreciate the opportunity provided by the referees to improve our manuscript. The text has been redrafted, new tables and a new figure have been added. Clean and trackchanged versions are supplied.
Listed below are the suggestions from each of the referees, with our responses in red text. The line numbering is cross-referenced to the new clean version of the manuscript.

Editor's comments
Clarify the scope (just UK or global?) and be more up-front about the (big) differences between the diseases.
We have now made clear in the title, abstract and the introduction that our intention is to look at similarities in the global management of these two diseases but using the UK responses to FMD in FMD 2001 and to COVID19 in 2020 as representative points of reference. We think that moving from the specific to the general is a good way to explore this topic.
One table (current Table S1) already showed similarities and differences in transmission characteristics between the viruses. We have added a section that compares the morbidity and mortality of FMD and COVID19 (supplementary text lines 31-48). We have added a new table to show similarities and differences with respect to control of FMD and COVID19 (Table S2).

Paper exceeds allowed length
As well as improving the conciseness of language, we have moved additional materials, including two tables to the supplementary materials in order to reduce the paper to 10 printed pages.

Referee #1
To have its greatest impact for a general reader this reviewer feels that it needs to be a much shorter and concise document.
Many of the referees' comments require the supply of additional information. To keep the paper as concise as possible, we have added some new materials including tables and moved these along with some of the original text to the supplementary materials.
The following are some suggestions 1. Table 1 is very effective. A number of Tables with comparisons between FMD and COVID19 under a number of different heading would greatly help the reader to understand the range of problem areas in the practice of One Health. A list of current data sources would also be helpful. Table 1 (now Table S1) has had some extra information added on R0 and the references have been strengthened. Two new tables and additional references have been added.
2. While the authors are very clear about the outcome of the inquiry into FMD2001, it is lost in the detail of the manuscript. The lessons learned, which can all be alineslied to COVID 19 should form a central theme of the manuscript and be included in the abstract.
A new Table (now Table 1) sets this out and provides references to the original reports. The key overall "one health" conclusion is now included in the abstract.
3. The similarities in the lack of test capacity must be highlighted. While COVID 19 is considered a new virus, it is related to other SARS virus so the lack of investment in the development of accurate test capacity was not learned from FMD but must be learned for the next animal or human pandemic Additional details and emphasis added -lines 158-174; 254-268; 327-330; Tables 1 and S2. 4. While the authors report the percentage of the total herd that was culled, some standardisation of the death rate such as the proportion of the population of England who died from COVID 19 rather than the actual numbers should be provided for a given time (April 30th 2020). Comparison of crude numbers, as currently presented in the media has little meaning.
Calculations reworked -lines 63-64. 5. A short section on the priorities for Public Health or One Health Preparedness, based on lessons learned from both Epidemics would also be helpful Included in abstract and conclusion -lines 22-24; 345-350 6. Table 2 is cited in the conclusion but I feel this is Table 1. New Table numbering Referee #2 1) The title should probably include reference to the "the UK" See new title in which this is addressed.
2) Please compare estimates of R0 for both diseases. Can you say anything about the severity of the diseases?
Both addressed - Table S1 and supplementary text lines 31-48 3) While COVID-19 spread rapidly in the UK following an exponential growth pattern, the 2001 FMD epidemics spread following a slower growth pattern in the number of premises infected (see https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6132414/) Now discussed -lines 188-190.

4) Please display the epidemic curves for both epidemics including a timeline of the key interventions
Done -see new Figure 2

Referee #3
However, there are gross differences that make the comparison less compelling: many of the disease characteristics are quite different (e.g. FMD affects multiple species, with different levels of susceptibility and transmission routes, the presence of carrier species and wild reservoirs), sources of cost in the disease (FMD rarely kills -the main cost is in the trade response it induces), and obvious differences in management (we cannot cull people). Many of these are left out of the paper.
These are now covered in greater detail (Tables S1 & S2, supplementary text, main text lines 32; 76; 296-308; Supplementary text lines 32-48), although it is not possible to keep the MS short and provide a textbook account of FMD. All of the specific points raised above (reservoirs, relative mortality, management differences such as culling) are now covered in text and/or supplementary tables and text.
Because the paper switches between a UK and global focus, it is unclear to me what the primary research question is. If the primary research question is "What can we learn by comparing the current pandemic to the 2001 FMD epidemic?" then the paper must be revised to encompass the broader management strategies and incentives of FMD and COVID and provide a more balanced comparison of the two diseases. I do not need to compare COVID19 to FMD to know that early detection, testing, and movement restrictions are needed prevent spread and limit infections. But that is not to say that there are interesting lessons to be learned from the two. While some of the important details need to be added to more fully describe the 2001 epidemic, there is a story about the initial seeding and effectiveness of management efforts in regions that were overwhelmed or not by the number of FMD cases. In addition, there are some interesting questions regarding the global management of FMD, and the trade offs associated with eradicating FMD in developing countries vs. welfare, that will be applicable to future COVID19 management We think it is possible to cover two main themes: 1) were the relevant lessons of FMD in 2001 applied to COVID19 in 2020, and 2) is international control of FMD relevant to future COVID19 management.
If the primary research question is "Can we use the 2001 FMD epidemic to evaluate if the United Kingdom improved its biosecurity and management policies for facing new epidemics?" then the paper is in better shape. That being said, much of the paper will need to be reworked and the authors will need to decide on when to focus on the UK versus the global story.
We believe that we have addressed this research question and have reworked the paper in multiple ways to make the distinction between UK and global controls clearer. For example, by making clearer some of the different approaches needed for FMD control in endemic versus FMD-free countries and regions. However, inevitable overlap remains between UK and wider responses to FMD and COVID19.
Regardless of the editorial decision, I believe that there need to be a number of changes to improve the manuscript. First, the scale of the study -particularly that of the 2001 epidemicneeds to be clarified. The authors flip back and forth between management within the United Kingdom and global management of FMD in 2001, which is confusing and changes what we can say about disease management. The World Organization of Animal Health maintains a set of five disease-free categorization ranging from disease-free, no vaccinations (low risk) to not disease-free (high risk). Countries with and without FMD face very different decisions in how they manage disease risk within their own countries but also on the global market. Questions of welfare and tradeoffs within developed and developing countries are important for FMD management (see the 2012 Global FMD Control Strategy) and will be for COVID19.In this same vein, the nature of disease-free designations, and how they impact the different management strategies is important. Understanding why people choose their management decisions and how they are incentivized or compensated (maybe too well for the 2001 FMD epidemic) matters for the current management of an epidemic but also for how they will view and react to future risks of disease.
We have reorganised the materials and headings and this makes it clearer where we are talking about the UK 2001 epidemic and wider control measures. Some further details are given of the international trade standards set for FMD by The World Organisation for Animal Health. The relevance of these measures to alternative future management options for COVID19 appears to have become more and not less compelling since we submitted our first draft.
The temporal scales of the epidemics are also important. Where do each of the "phases" in the manuscript fall on the epidemic curves for each? Could the authors include a figure of each epidemic labeling the phases? There are some inconsistencies in the timing between the phases, such as testing centers being overwhelmed (which is in phase 1 but could easily fall in phase 2 and 3, no?).
We have introduced a figure showing the epidemic curves and main events for each situation.
Finally, if we are going to compare the two diseases, then it should be done so in a complete fashion. Many of the reasons that FMD is such a pain to manage are mostly ignored. Specifically, FMD infects virtually every cloven hoofed animal in the world, can be spread by multiple pathways with different animals exhibiting different susceptibilities and pathways of viral shedding, the presence of carrier animals, and wild disease reservoirs. While I understand that this will cause the comparison to diverge, it is important to remain transparent and open with the science of the two diseases.
Not all knowledge about the biology and control of FMD can be summarised in this article. Nevertheless, we have included additional details about its host range, morbidity and mortality (in the supplementary materials) and about international FMD controls. I am including a number of other comments and suggestions embedded in the attached .pdf file. Please let me know if they are not viewable.