One of the sleepers in population dynamics has always been the role of disease in population limitation and population fluctuations. Part of the reason for this is that disease studies need cooperation between skilled ecologists and skilled microbiologists. Another problem is the possibility of infinite regress in looking for disease agents as a cause of population change in natural populations – e.g. if it is not virus X, there are hundreds of other viruses that might be the culprit. In both North America and Europe one focus of concern has been on the hantavirus group (Luis et al. 2010; Mills et al. 2010, Davis et al. 2005, Mills et al. 1999). Hantaviruses come in many different forms and are typically carried by rodent species. Some varieties produce hemorrhagic fever with renal syndrome in Europe, Asia and Africa, but in the Americas the main disease of concern is HPS (hantavirus pulmonary syndrome). It is no surprise that often emerging diseases are studied only because some humans die from them. As of 2016, 690 cases of hantavirus pulmonary syndrome have been recorded in the USA, and 36% of these cases resulted in death. The reverse question of what the disease is doing to the animal population gets rather less attention typically than the human disease problem. The example I want to discuss here is the Sin Nombre virus (SNV) in deer mice (Peromyscus spp.), widespread rodents in North America.

The hantavirus outbreak in the Southwestern USA in the 1990s caused numerous human deaths and produced a number of field studies that showed a patchy pattern of infection among deer mice in Arizona and Colorado (Mills et al. 1999). Male mice were infected more than females and the suggestion was that males fighting for territories were infecting one another directly when population densities were high. The call for long-term studies went out and several studies from 3-5 years were carried out in the late 1990s until the problem of infection in the human population became less of an issue compared with other diseases such as Ebola in other parts of the world. The shift in concern resulted in reduced funding for field studies in North America.

In 1994 Rick Douglass and his research team began long term studies on the Sin Nombre virus in deer mice using 18 live trapping areas of 1 ha each spread across Montana and placed in a variety of habitats (Douglass et al. 2001). Long-term for their study was 15 years, all this at a time when 2-3 year studies were thought to be sufficient to unravel the nexus of infection and transmission. The idea was to complement in Montana similar rodent research in Arizona, New Mexico, and Colorado. The results are fascinating and important because they illustrate the importance of long term research and the understanding of what a well designed field study can produce.

Rightfully many of the hantavirus studies were focused on the human connection, but what I want to emphasize here is the impact of this virus on the rodent populations. Luis et al. (2012) estimated that male Peromyscus had their monthly survival rate reduced from 0.67 to 0.58 if they were seropositive, a 13% reduction, but females showed no effect of hantavirus on survival so that infected and uninfected females survived equally well. Hantavirus does reduce body growth rates of infected male mice. One consequence of these findings should be that the growth rate of Peromyscus populations in Montana should be only slightly affected by hantavirus infections, since it is the female component of the population that drives numbers. There are limitations to these conclusions since juveniles too young to live trap could suffer mortality that at present cannot be measured. The threshold for hantavirus transmission in these Peromyscus populations was about 17 individuals per ha (Luis et al. 2015), implying that hantavirus would disappear in populations smaller than this because it would not transmit. The consequence for us is that human hantavirus infections in North America are much more likely when deer mouse populations are high, and by monitoring deer mice ecologists can broadcast warnings when there are increased possibilities of infection with this lethal disease.

The details about the Sin Nombre hantavirus in North America are well covered in these and other papers. The most important general message from this research has been the need for long term studies to get at what might initially seem to be a simple population problem (Carver et al. 2015). There are a host of other viruses that infect rodent species and many other mammals and birds about which we know very little. The path to understanding the effects of these viruses on the animals they infect and their potential for human transmission will require much detailed work over a longer time period than what is now the funding horizon of our granting agencies. The Montana studies on the Sin Nombre virus required ecologists to trap for 20 years with more than 851,000 trap nights to catch 16,608 deer mice, and collect 10,572 blood samples to assess infections and gain an understanding of this virus disease. The problem too often is that it is easy to find ecologists and virologists keen to cooperate in these studies of disease, but it is not easy to find the long term funding that looks at these ecological problems in the time scale of 10-20 years or more. We need much more long term thinking about ecological problems and the funding to support team efforts on difficult problems that are not soluble in a 3-year time frame.

Carver, S., Mills, J.N., Parmenter, C.A., Parmenter, R.R., Richardson, K.S., Harris, R.L., Douglass, R.J., Kuenzi, A.J., and Luis, A.D. 2015. Toward a mechanistic understanding of environmentally forced zoonotic disease emergence: Sin Nombre hantavirus. BioScience 65(7): 651-666. doi: 10.1093/biosci/biv047.

Davis, S., Calvet, E., and Leirs, H. 2005. Fluctuating rodent populations and risk to humans from rodent-borne zoonoses. Vector-Borne and Zoonotic Diseases 5(4): 305-314.

Douglass, R.J., Wilson, T., Semmens, W.J., Zanto, S.N., Bond, C.W., Van Horn, R.C., and Mills, J.N. 2001. Longitudinal studies of Sin Nombre virus in deer mouse-dominated ecosystems of Montana. American Journal of Tropical Medicine and Hygiene 65(1): 33-41.

Luis, A.D., Douglass, R.J., Hudson, P.J., Mills, J.N., and Bjørnstad, O.N. 2012. Sin Nombre hantavirus decreases survival of male deer mice. Oecologia 169(2): 431-439. doi: 10.1007/s00442-011-2219-2.

Luis, A.D., Douglass, R.J., Mills, J.N., and Bjørnstad, O.N. 2010. The effect of seasonality, density and climate on the population dynamics of Montana deer mice, important reservoir hosts for Sin Nombre hantavirus. Journal of Animal Ecology 79(2): 462-470. doi: 10.1111/j.1365-2656.2009.01646.x.

Luis, A.D., Douglass, R.J., Mills, J.N., and Bjørnstad, O.N. 2015. Environmental fluctuations lead to predictability in Sin Nombre hantavirus outbreaks. Ecology 96(6): 1691-1701. doi: 10.1890/14-1910.1.

Mills, J.N., Amman, B.R., and Glass, G.E. 2010. Ecology of hantaviruses and their hosts in North America. Vector-Borne and Zoonotic Diseases 10(6): 563-574. doi: 10.1089/vbz.2009.0018.

Mills, J.N., Ksiazek, T.G., Peters, C.J., and Childs, J.E. 1999. Long-term studies of hantavirus reservoir populations in the southwestern United States: a synthesis. Emerging Infectious Diseases 5(1): 135-142.