Journal of Animal
Ecology 2007
76, 711–721
© 2007 The Authors.
Journal compilation
© 2007 British
Ecological Society
Blackwell Publishing Ltd
Quantifying the disease transmission function: effects of
density on Batrachochytrium dendrobatidis transmission in
the mountain yellow-legged frog Rana muscosa
LARA J. RACHOWICZ*† and CHERYL J. BRIGGS*
*Department of Integrative Biology, University of California, Berkeley, CA 94720-3140, USA; and †Resources
Management and Sciences, Yosemite National Park, 5083 Foresta Road, PO Box 700, El Portal, CA 95318, USA
Summary
1. Chytridiomycosis is an emerging infectious disease of amphibians, caused by the
fungal pathogen Batrachochytrium dendrobatidis, which has been implicated recently in
population declines and possible extinctions throughout the world.
2. The transmission rate of this pathogen was quantified in the mountain yellow-legged
frog Rana muscosa through laboratory and field experiments, and a maximum likelihood
approach was used to determine the form of the transmission function that was best
supported by the experimental data.
3. The proportion of R. muscosa tadpole hosts that became infected increased with the
number of previously infected R. muscosa tadpoles to which they were exposed, as
would be expected in an infectious disease.
4. The laboratory experiment revealed some support for a transmission function in
which the transmission rate levels off as the density of infected individuals increases.
However, there was not enough power to distinguish between a frequency-dependent
form and several other asymptotic forms of the transmission function.
5. The impacts of crowding and temperature on transmission were also investigated;
however, neither of these factors significantly affected the transmission rate.
Key-words: amphibian decline, disease transmission, extinction, host–pathogen interaction,
Sierra Nevada.
Journal of Animal Ecology (2007) 76, 711–721
doi: 10.1111/j.1365-2656.2007.01256.x
Introduction
Chytridiomycosis, an emerging infectious disease of
amphibians caused by the chytrid fungal pathogen
Batrachochytrium dendrobatidis (Berger et al . 1998;
Longcore, Pessier & Nichols 1999), has been implicated
in the dramatic population declines, and even complete
extinctions, of numerous amphibian species throughout
the world (Daszak, Cunningham & Hyatt 2003). In
mountain yellow-legged frogs Rana muscosa Camp, B.
dendrobatidis has been associated with the extinction of
hundreds of local populations in the Sierra Nevada
mountains of California in recent years (Rachowicz
et al . 2006).
Although the potential for disease to impact, or even
regulate, host population dynamics has long been
recognized (Anderson & May 1992; Hudson et al .
2002), simple disease models predict that disease on its
own cannot cause the complete extinction of a popu-
lation or species (Anderson & May 1992; de Castro &
Bolker 2005). The reason for this is that simple models
of directly transmitted pathogens generally assume
that disease transmission takes on a density-dependent
(mass action) form. Density-dependent disease trans-
mission leads to host density thresholds below which
the disease cannot invade or persist (McCallum,
Barlow & Hone 2001; Lloyd-Smith et al . 2005), such
that the disease is predicted to die out before the host
population drops to extinction. However, if pathogen
transmission takes on a frequency-dependent, rather
than a density-dependent form, disease-induced extinction
of the host is possible because frequency-dependent
transmission results in no such host density threshold
(Getz & Pickering 1983; McCallum et al . 2001). There-
fore, quantification of the disease transmission process
is extremely important for understanding the potential
Correspondence: L. J. Rachowicz, Department of Integrative
Biology, University of California, Berkeley, CA 94720-3140,
USA. E-mail: lara.rachowicz@nps.gov