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Author: Hilary Stern[a][i]

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  1. ↑ Animal Hospital of Soquel, Santa Cruz CA USA
  1. ↑ hsterndvm@gmail.com

This article has been withdrawn by the submitting author for publication by the WikiJournal of Science.

It is archived here as a record. Discussion can be viewed below.

Plagiarism check

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Latest comment: 5 years ago1 comment1 person in discussion

  Pass. Report from WMF copyvios tool. Whole article appears in other site that explicitly licenses cc by-sa). Some trivial overlap observed more broadly with common phrases (e.g. "a safe and effective myxomatosis vaccine for pet rabbits") but not considered plagiarism. No genuine plagiarism detected. T.Shafee(Evo﹠Evo)talk 01:36, 21 August 2019 (UTC)Reply

First peer review

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reviewer-annotated pdf

Review by Morgan P Kain , Stanford
These assessment comments were submitted on , and refer to this previous version of the article

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Second peer review

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reviewer-annotated pdf

Review by Justine Philip , University of New England, Australia; Museam Victoria, Australia
These assessment comments were submitted on , and refer to this previous version of the article

The paper covers the scientific description of the Myxomatosis virus and its presentation, however it is lacking in an account of the complex social, political and ethical impacts that this virus had on the scientific and general community. Myxomatosis exposed the dangers of employing biological warfare against "pest" species by the agrichemical farming industry in peacetime. The term vermin or pest species, is in itself a cultural construct, not a scientific description; rabbits might have been problematic to sheep farmers but were a valuable resource to other members of the community, and a valued cultural asset in their country of origin. The Myxomatosis virus was never released, it escaped escaped confinement and then it proved impossible for governments to police the movement of the virus once it was out in the community (Olsen 1998, Cooke 2014).

The resulting impact was removal of a valuable resource that while not a huge export earner, was a valuable and free resource, particularly important to low income and indigenous communities (Philip 2017, Philip 2018). These communities were deprived of a plentiful, nutritious food source, and often a source of primary or supplementary income, supplying industries reliant on rabbit fur, pelts, meat etc. These industries went bankrupt after the release of Myxomatosis, and trappers and hunters were superseded by a new wave of increasingly intensive agricultural farming operations. By the 1960s, it was cheaper to raise a chicken from a hatchling to slaughter in a factory farm, than to go out and shoot a wild rabbit - chickens replaced rabbit meat as a staple food source for much of the population (see Ether and Cottle, 2015).

A thorough account of Myxomatosis also must include content on pertinent issues such ethical considerations, and the extreme animal cruelty and suffering that resulted from the disease. There was widespread condemnation of this use of biological warfare against wildlife by leading animal welfare organisations, veterinarians, and public figures (Winston Churchill for one) and general public. The negative impacts on ecosystem health also are often absent from scientific studies, including the possible extinction of symbiotic species.

The exposure of the community to millions of infected and dying animals was deeply distressing to children to witness, and concerned many elder members of the community. Animal protection groups worldwide condemned the release of the virus. The RSPCA in the United Kingdom raised significant funds to help euthanise as many rabbits as possible, to prevent their inevitable lengthy and painful death. The disease left millions of rabbits blinded and starving, an estimated 99% of rabbits in the United kingdom were killed by the disease, and 90-90% in France following it's illegal release in Europe.

Important areas to cover

The impossibility of containment

A key point with myxomatosis is that it did not behave the way the scientific trials in Australia in 1949 had predicted that it would. After extensive trials in the late 1940s, they had concluded that an introduction would be a failure and were about to abandon the study, when the myxomatosis virus escaped from a field site in the Riverina of NSW (see PERSPECTIVE Promising new weapon in the war on rabbits 1992). With unseasonably favourable weather for the mosquito vectors that year, the results were staggering - myxomatosis decimated the Australian rabbit population (Cooke, 2014 p.58). This highlights the unpredictability and danger of employing biological weapons, a lesson repeated again with the introduction of calicivirus in 1995 that also escaped during trials and spread fast across Australia (Olsen, 1998, pp. 65-66). Cooke (2014 p. 65) wrote that the virus did not behave "as predicted based on European experience."

The impact on the European and UK rabbit populations.

News of Australia's success in reducing rabbit numbers quickly reached Europe. All it took was one elderly, errant, self motivated physician. Dr Paul Delille took it upon himself to obtain a virus sample via his scientific contacts, and he released it on his farm near Deux in France in 1952 (Bartrip 2009). By August 1953 the virus had spread through most of France and many other Continental countries before jumping the English Channel and spreading like wildfire through the UK - aided by many farmers keen to rid their lands of "the rabbit pest".

Animal cruelty and welfare considerations.

The RSPCA published advertisements in the UK press:

"Stop the deliberate spreading of MYXOMATOSIS!—Victims of this horrible disease—blind, misshapen, tormented—are being caught for sale as carriers, to be let free in infection-free areas ... . Nothing can justify this callous encouragement of animal suffering, and the RSPCA appeals for your moral and material support in demanding an immediate legal ban."

From the 1890s through to the end of World War 2, rabbits provided a thriving trade in meat and fur for Australia. The industry prospered through times of drought, war, and economic downturns – events that severely impacted on the sheep and cattle industries and resulted in decades of hardship for pastoralists (Eather & Cottle, 2015). At the time that the rabbit industry was first expanding, the national sheep flock decreased from 106 million to 54 million (1892 to 1904). “In 1929 the rabbit industry was reported to be Australia’s largest employer of labor” (Eather & Cottle, 2015, p. 1).

At this time, over 20,000 trappers worked full-time trapping for carcasses or skins, or poisoning for skins. Thousands were employed in numerous freezer works located in rural towns and capital cities; grading, sorting, packing, skinning and transporting carcasses by the tens of millions. In addition there were thousands employed in the fur industry, and selling rabbit meat directly to the public through street stalls and shops, making felt hats out of the rabbit skins (Eather & Cottle, 2015). Philip, 2017, recorded:

Even the ‘scraps’ went into fertilizer, animal feed, and to make gelatin. Around four billion rabbit skins were exported between 1904 and 1947. An estimated 27 million rabbits were consumed by Australians each year during the 1940s (ABC, 2015). Australian soldiers in World War 2 marched into battle wearing slouch hats made of rabbit skins – ten rabbits per hat, and Australia produced 5,500,000 hats during the war.

Last of the Lantern Swingers. A story of the rabbit industry in Sunraysia, historian G B Eggleton concluded that “the industry was a far better solution to the rabbit problem than either poisoning or myxomatosis.” The Governments of Australia and New Zealand banned the trade in rabbit products in the 1950s bringing an end to what had been a thriving economy. The impact of rabbits on the environment was arguably no more damaging than the sheep and cattle industry.

An example of public misinformation about the rabbit pest appears here with the publication of a photo in 1905 claiming that rabbits had killed everything outside of the fenceline. Environmental historian Don Garden (2005), featured the 1905 photo, Figure 16-9, of rabbit-proof fencing, in the Australia, New Zealand, and the Pacific : an Environmental History, as an illustration of the propaganda supporting rabbit eradication in the early days of Federation. Garden (2005, p. 74) recorded:

This image was intended to demonstrate the effectiveness of rabbit- proof netting fences by showing the contrast between a paddock eaten bare by rabbits and adjacent area protected from them. However, the picture may not be so clear. Desperate rabbits might well be able to cross such a low fence, so one suspects that overgrazing by sheep was part of the reason for the loss of vegetation.

[Image in attached pdf] Figure 16-9 “The rabbit pest: two sides of a netting fence.” Cobar, New South Wales 1905. Source: State Library of New South Wales At Work and Play 02766

To end, consider the quote from the Secretary of the World League for Protection of Animals of biological warfare in 1951: "no good ever came out of cruelty."

References

Third peer review

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Review by anonymous peer reviewer ,
These assessment comments were submitted on , and refer to this previous version of the article

This is a good summary by Hilary et al about myxomatosis. While it is well developed in some sections, others require more detail, especially taking into account the broader importance of myxomatosis in the context of evolutionary biology. I am not qualified to comment on the clinical aspects of myxomatosis, so my comments are not addressing those sections. My comments to each section are below:

Section: Summary

  • “The virus was intentionally introduced in Australia, France, and Chile in the 1950s to control wild European rabbit populations.” The circumstances of the virus release as well as the attitude towards rabbits were extremely different in Australia and France. I am slightly concerned that the word “intentional” in this phrase does not reflect the important distinction between the governmental decision of releasing the virus in Australia, versus the isolated act of a single individual in France.
  • In addition to the species jump, the summary could also emphasise that a major reason why myxomatosis is so well studied is that it is an extraordinary example of co-evolution in the wild (see comments below)

Section: Cause

  • “Different strains exist which vary in their virulence.” This phrase could be more developed. In particular, the method used to grade the virus strains (i.e. infecting rabbits with different strains and evaluating the clinical signs and survival rates). It was ultimately this phenotypical grading that allowed to track both the evolution of myxoma virus virulence in the wild in the years following introduction. Moreover, it was the fact that there was an original strain in the lab that conversely allowed to assess the evolution of resistance in wild rabbits. For a summary see Kerr, Peter J. ""Myxomatosis in Australia and Europe: a model for emerging infectious diseases."" Antiviral research 93.3 (2012): 387-415).
  • “While wild rabbits in Europe and Australia have developed some immunity to the virus”. It would be good to clarify what it is meant by immunity (acquired immunity?). Rabbits also evolved genetic resistance (see below)

Section: Use as a population control agent

  • This section would benefit from some context on the motivation to use of myxomatosis as biological control. In particular, the ecological/economic impact and the dimension of the rabbit problem in Australia and some of the initial unsuccessful attempts to control it, such as the large fences. It should also reflect the extensive research and trials made in the earlier decades to evaluate the impact and potential success of myxomatosis. This will allow putting in context the use and the implementation of such a drastic method of population control. This book provides extensive insight on this matter: “Fenner, F. & Fantini, B., 1999. Biological Control of Vertebrate Pests: The History of Myxomatosis - an Experiment in Evolution, New York, NY, USA: CABI publishing.).
  • I am not sure how well this fit with the article but some comment on the ethical aspects of the release of myxomatosis and the conservation paradox of rabbits (ecologically important and endangered in the native range; pests in many introduced locations) could be interesting to stem new discussions.
  • “The long-term failure of this strategy has been due to natural selective pressures”. I would remove this phrase. It is debatable whether the use of myxomatosis was a failure and such interpretation can be often an oversimplification of the outcome. Myxoma virus continues to evolve and circulate in rabbit populations and while it is true that in many locations rabbits recovered, in many others the number of rabbits has never reached pre-myxomatosis numbers. This drop was also accompanied by a positive impact on the ecology of the affected areas. I would recommend reading the section “5.3. Success or failure of biological control?” from Kerr, Peter J. ""Myxomatosis in Australia and Europe: a model for emerging infectious diseases."" Antiviral research 93.3 (2012): 387-415).) where he addresses precisely this point. See also: Sumption, K. J., and J. R. Flowerdew. ""The ecological effects of the decline in rabbits (Oryctolagus cuniculus L.) due to myxomatosis."" Mammal Review 15.4 (1985): 151-186.
  • In the Australia section, it might be worth mentioning that while rabbits brought to Australia in 1788, it is believed that the major source of the infestation occurred in 1859 after a release of wild rabbits in Victoria. It might be worth cross-referencing the putative responsible for this release (https://en.wikipedia.org/wiki/Thomas_Austin). See: “Fenner, F. & Fantini, B., 1999. Biological Control of Vertebrate Pests: The History of Myxomatosis - an Experiment in Evolution, New York, NY, USA: CABI publishing.).
  • In the Europe section, it might be worth to cross-reference Armand-Delille, the person responsible for the introduction of rabbits in France (https://en.wikipedia.org/wiki/Paul-F%C3%A9lix_Armand-Delille). It might also be worth to dedicate a small section highlighting the UK, as this was one of the countries where the impact of myxomatosis on rabbit populations was extensively studied and there is substantial literature (See articles from Ross, J. & Sanders).
  • “Unlike in Australia, however, strenuous efforts were made to stop the spread in Europe”. It could also be emphasised how this was due to the very distinct attitude towards rabbits in Australia versus Europe. In particular, in France, rabbits were quite an important species for hunting and there was a large rabbit-breeding industry. On the other hand, this clashed with farmers that saw them as pests (for more info see: “Fenner, F. & Fantini, B., 1999. Biological Control of Vertebrate Pests: The History of Myxomatosis - an Experiment in Evolution, New York, NY, USA: CABI publishing.).
  • “Some dissemination was clearly deliberate, such as the introduction into Britain in 1953” I have no access to the book referenced here but in all literature, I came across it has always been mentioned that the means of entry of myxomatosis in England is unknown (see Fenner, F. & Ratcliffe, F.N., 1965. Myxomatosis, Cambridge; New York: Cambridge University Press.)

Section: Use as an evolutionary model

  • I feel like this is the section that requires substantially more development. The coevolutionary arms-race between rabbits and myxomatosis has become a textbook example of host-parasite coevolution contributing to the development of conceptual and mathematical models of infectious disease evolution, pathogen evolution and optimal virulence (see: Anderson, R.M. & May, R.M., 1982. Coevolution of hosts and parasites. Parasitology, 85 (Pt 2), pp.411–426.; Dwyer, G., Levin, S.A. & Buttel, L., 1990. A simulation model of the population dynamics and evolution of myxomatosis. Ecological Monographs, pp.423–447; Saunders, I.W., 1980. A model for myxomatosis. Mathematical Biosciences, 48(1-2), pp.1–15.; Alizon, S. & Michalakis, Y., 2015. Adaptive virulence evolution: the good old fitness-based approach. Trends in Ecology & Evolution.)
  • Over the last decade multiple studies have also looked into the genetic changes associated to the virulence of the different strains showing that there are likely multiple genes involved and potentially different pathways to the same phenotype (e.g. Kerr, P.J. et al., 2012. Evolutionary history and attenuation of myxoma virus on two continents. PLoS pathogens, 8(10), p.e1002950.). Recently it has also been shown that the continuous co-evolution of these two organisms resulted, as predicted, in myxoma virus strains that are more virulent than the strain that caused the original epidemics (Kerr, Peter J., et al. ""Next step in the ongoing arms race between myxoma virus and wild rabbits in Australia is a novel disease phenotype."" Proceedings of the National Academy of Sciences 114.35 (2017): 9397-9402.
  • On the rabbit side of things, there is also extensive literature that discussed the independent emergence of genetic resistance in multiple locations (for a summary see: Kerr, Peter J. ""Myxomatosis in Australia and Europe: a model for emerging infectious diseases."" Antiviral research 93.3 (2012): 387-415). Recently it was also showed that this genetic resistance is the result in of parallel genetic changes in the same genes across the rabbit populations of Australia and Europe (Alves, Joel M., et al. ""Parallel adaptation of rabbit populations to myxoma virus."" Science 363.6433 (2019): 1319-1326.). The following perspective discusses some of the general implications of this finding (https://science.sciencemag.org/content/363/6433/1277).
  • It is also mentioned in the summary that myxomatosis is very well studied due to the species jump, but there is no additional info in this section to support this. On this topic, it was recently reported the jump of myxomatosis to hares which might have a place in some section of this article (see: Dalton, Kevin P., et al. ""Myxoma virus jumps species to the Iberian hare."" Transboundary and emerging diseases (2019))
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