Abstract

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.

Highlights

  • The extent that top-down forces from predators, bottom-up forces from resources, and abiotic forces such as weather interact to affect population growth in prey species is one of the most central questions in ecology [1,2,3]

  • Prior to the reintroduction of wolves to the Greater Yellowstone Ecosystem (GYE) in 1995 and 1996, over five decades of research and monitoring of Yellowstone elk dynamics consistently found that (1) variation in juvenile recruitment explained most of the annual variation in elk population growth and that (2) negative density dependence and winter severity were the strongest drivers of annual variation in recruitment [7,8,9,10,11]

  • In the GYE elk numbers declined by .50% after wolf reintroduction in some areas [15,16], exceeding the 5–30% declines predicted by models of wolf-elk dynamics [17]

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Summary

Introduction

The extent that top-down forces from predators, bottom-up forces from resources, and abiotic forces such as weather interact to affect population growth in prey species is one of the most central questions in ecology [1,2,3]. Prior to the reintroduction of wolves to the GYE in 1995 and 1996, over five decades of research and monitoring of Yellowstone elk dynamics consistently found that (1) variation in juvenile recruitment explained most of the annual variation in elk population growth and that (2) negative density dependence and winter severity were the strongest drivers of annual variation in recruitment [7,8,9,10,11] These patterns are commonly seen in other ecosystems, and in other long-lived iteroparous species [12,13,14]. In the GYE elk numbers declined by .50% after wolf reintroduction in some areas [15,16], exceeding the 5–30% declines predicted by models of wolf-elk dynamics [17] This discrepancy prompted more recent analyses of GYE elk population dynamics that have incorporated density dependence and winter severity, and considered recent data on predation rates by wolves, other climate variables, and harvest rates from elk hunting. Offtake from hunters, of reproductive females, exceeded offtake by wolves in the years immediately following reintroduction and hunting has been suggested as a factor possibly explaining declining elk abundance following wolf reintroduction [15,18,19,23]

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