Plant community succession following ungulate exclusion in a temperate rainforest

Ecosystem structure and processes of coastal temperate rainforests of the Pacific Northwest are thought to be strongly influenced by herbivory primarily of Roosevelt elk (Cervus elaphus roosevelti) and secondarily of Columbian black-tailed deer (Odocoileus hemionus columbianus). Two large (0.5-ha) exclosures were built in old-growth coniferous rainforest communities in Olympic National Park, Washington, during 1979 to study these effects. Cover of shrubs, ferns, herbs, and graminoids and numbers of tree seedlings were described over 36 yr. Results show a sequence following ungulate exclusion of early release of shrubs, ferns, and herbs followed by eventual dominance of shrubs as other vegetation layers become shaded. Short-term responses of individual species reflected functional traits related to ability to avoid or tolerate herbivory. Over the longer term, effects reflected changing competitive relationships among vegetation layers and other ecosystem dynamics such as the provision of fallen trees in the appropriate decay class to serve as establishment substrate for tree seedlings. In aggregate, vegetation composition shifted after 36 yr from a system dominated by herbaceous cover with a major graminoid component to one dominated by shrubs (5- to 6-fold absolute increase) and ferns (5–7% increase in absolute cover), less absolute herb cover (15–20% loss), and almost no graminoids (