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it was used by many managers as well as regional high school and university
students. Some of the predictions gleaned from the model were incorporated
directly into the wolf recovery program. For example, stochastic population
projections revealed that the small number of wolves proposed for releases
were likely to go extinct within a year or two, but that the success of wolf
releases could be enhanced considerably by scheduling repeated releases
(Boyce 1992b). Indeed, this is exactly what the government did, and appar-
ently the program has been highly successful (Fritts et al. 1995).
Conclusion
j
Seventy years ago, when Lotka (1925) and Volterra (1926) first introduced
predator-prey models, a debate raged about whether mathematical formula-
tions could be trusted to provide insight into real-life ecological systems
(Kingsland 1985). During the golden era of rapid theoretical development in
ecology and population genetics of the 1920s and 1930s, some feared that bio-
logical facts would give way to mathematical theory (Thompson 1937).
Amazingly, the same battle between modelers and empiricists still rages today.
Field ecologists and managers still make disparaging remarks about models
that they do not understand (Mech 1995; Schullery 1995) while theoretical
ecologists continue to explore the dynamics of models for which we have yet
to find clear examples in nature (Schaffer 1988).
I hold the basic premise of Galileo that the “Book of Nature is written in
mathematical form.” Like Galileo I believe that our understanding of nature
(read predator-prey dynamics) will proceed only through a rigorous iteration
of mathematical models with “cimento” (experiments). We have an extensive
body of literature on predator-prey models. We also have an extensive body of
literature on the behavior and ecology of predators and prey (Gittleman 1989,
1996). However, empirical studies of predators and prey seem preoccupied
with techniques and are seldom placed in the context of the theory. Only when
the theory and the fieldwork become integrated will we begin to develop a sci-
ence on the level of sophistication that Galileo envisioned 400 years ago.
Acknowledgments
Thanks to Todd Fuller and Luigi Boitani for inviting my participation in the conference
and to Danilo Mainardi for support from the International School of Ethology. I benefited
from suggestions and discussions with Gary White, Todd Fuller, Charles Krebs, Joe Elkin-
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