Contributions of forage competition, harvest, and climate fluctuation to changes in population growth of northern white-tailed deer

Recently there has been considerable interest in determining the relative roles of endogenous (density-dependent) and exogenous (density-independent) factors in driving the population dynamics of free-ranging ungulates. We used time-series analysis to estimate the relative contributions of density-dependent forage competition, climatic fluctuation, and harvesting on the population dynamics of white-tailed deer (Odocoileus virginianus) in Nova Scotia, Canada, from 1983 to 2000. A model incorporating the population density 2 years previous, an interaction term for the harvest of females and population density 2 years previous, and the total snowfall during the previous 2 winters explained 80% of the variation in inter-annual population growth rate. Natality of adult females was negatively related to deer density during the present winter, whereas that of yearlings may have been correlated with the snowfall of three winters previous. Natality of fawns was related to deer density and total snowfall during the previous winter. Coyotes (Canis latrans) prey extensively on deer fawns in northeastern North America and the annual harvest of snowshoe hares (Lepus americanus), the major alternate prey of coyotes, explained 48% of the inter-annual variation in fawn recruitment. The proportions of fawn, yearling, and adult deer suffering from severe malnutrition during late winter were all correlated with deer density during the present winter. We conclude that the limiting effects of winter weather on over-winter survival of deer may be cumulative over two consecutive winters. During the late 1980s, density dependence and winter severity acted in concert to effect substantial declines in deer population growth both by effecting winter losses directly and by exacerbating predation by coyotes. During this period liberal harvesting did not relieve density-dependent forage competition and probably accelerated the decline.