Excavation of red squirrel middens by grizzly bears in the whitebark pine zone

Journal of Applied Ecology
By:  and 



  1. Whitebark pine seeds Pinus albicaulis are an important food of grizzly Ursus arctos horribilis bears wherever whitebark pine is abundant in the contiguous United States of America; availability of seeds affects the distribution of bears, and the level of conflict between bears and humans. Almost all of the seeds consumed by bears are excavated from middens where red squirrels Tamiasciurus hudsonicus have cached whitebark pine cones.
  2. Relationships among the occupancy of middens by squirrels, the excavation of middens by bears, and site features were investigated in this study. Data were collected from radio-marked bears and from middens located from line transects on two study sites in the Yellowstone ecosystem.
  3. Densities of active middens were positively related to lodgepole pine Pinus contorta basal area and negatively related to steepness of slope.
  4. The probability that a midden was occupied by a squirrel (i.e. active) was positively related to lodgepole pine basal area in the surrounding stand, size of the midden and size of the whitebark pine cone crop, and negatively related to elevation and to bear excavation during the previous 2-12 months.
  5. The probability that a midden had been excavated by a bear during the previous 12 months was positively related to size of the midden, and to whitebark pine basal area and cone crop, and negatively related to nearness of roads and town sites.
  6. The influence of midden size on bear use was attributable to a positive relationship with the number of excavated cones. The positive association between bear excavations and whitebark pine basal area or cone crops was attributable to availability of pine seeds.
  7. Grizzly bears would benefit from the minimization of roads and other human facilities in the whitebark pine zone and from increases in the availability of whitebark pine seeds, potentially achieved by increasing the numbers of cone-producing whitebark pine trees, especially in lower elevations of the whitebark pine zone where red squirrels are more abundant.

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Additional publication details

Publication type Article
Publication Subtype Journal Article
Title Excavation of red squirrel middens by grizzly bears in the whitebark pine zone
Series title Journal of Applied Ecology
DOI 10.2307/2405283
Volume 34
Issue 4
Year Published 1997
Language English
Publisher British Ecological Society
Contributing office(s) Forest and Rangeland Ecosystem Science Center
Description 15 p.
First page 926
Last page 940
Country United States
Online Only (Y/N) N
Additional Online Files (Y/N) N
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