Trophic Cascades: Wolf Reintroduction and Ecosystem Restoration in Yellowstone
Elk Behavioral Changes Due to Wolves
- Elk historically defend themselves from wolves by early detection, which is difficult in dense shrub areas like river bottoms where cottonwood and willows grow.
- The dense cover in river bottoms reduces clear lines of sight, making these areas dangerous for elk as wolves can ambush them.
- Consequently, elk avoid spending time in willow thickets by streams when wolves are present.
Ecological Consequences: Trophic Cascade in Yellowstone
- Absence of Wolves: Prior to wolf reintroduction, elk spent significant time in cool, shady river bottom areas due to abundant food (cottonwood, willows) and fresh water. This led to overbrowsing of riparian vegetation.
- Wolf Reintroduction: The reintroduction of wolves altered elk behavior, making river bottoms dangerous. Elk reduced their time in these areas.
- This behavioral change by elk, driven by wolf presence, led to a trophic cascade, restoring vegetation along stream banks.
Observational Evidence in Yellowstone National Park
- Early Observations: Even in the first few years after wolf reintroduction (when numbers were low), scientists anticipated and began documenting changes, suggesting a pre-existing understanding of wolf effects on ecosystems.
- Initial Baseline (e.g., Spring 1996): Researchers took photographs of stream banks, observing high water due to melting snow and a notable lack of vegetation along the riverbanks, likely due to historical elk browsing.
- Later Observations (e.g., Spring 2002): Approximately 6 years later, after wolves had been present for 5 to 7 years, repeat photographs showed significant changes, with willows beginning to grow robustly along the stream banks.
- This recovery of vegetation was taken as strong evidence of a trophic cascade: wolves altered elk behavior, which in turn reduced browsing pressure, allowing riparian vegetation to recover.
Quantitative Data and Further Confirmation (Veshe and Ripple, 2007)
- Wolf Population Timeline: Data presented by Veshe and Ripple provides a timeline of wolf reintroduction and population growth:
- 1992-1995: Yellowstone had 0 wolves.
- 1996: Approximately 20 wolves were reintroduced in an initial batch.
- The wolf population increased steadily thereafter.
- Aspen Height as an Indicator: Aspen trees are another preferred food source for elk. The height of an aspen tree is a key indicator of elk browsing activity.
- An aspen tree shorter than an elk's maximum reach is considered "in trouble" because elk can and will browse it heavily.
- Measuring the average height of aspen trees allows researchers to assess the intensity of elk browsing.
- Survey Methodology: Researchers conducted transects in various parts of the park, surveying aspen tree heights in both riparian (streamside) and upland settings during the late 1990s.
- Findings: The data confirmed that as wolf numbers increased, aspen trees experienced reduced browsing and were able to grow taller, especially in areas where elk previously concentrated.
- Wolf Population Timeline: Data presented by Veshe and Ripple provides a timeline of wolf reintroduction and population growth:
Generalizability of Trophic Cascades
- The observed patterns in Yellowstone are not an isolated incident.
- Similar ecological patterns are repeated in other national parks across the country where large predators have been reintroduced.
- Example: In Olympic National Park, the reintroduction of mountain lions has led to similar changes in prey behavior and subsequent forest recovery.
- This consistent pattern across different ecosystems (involving "large predators, their prey, and forests") provides compelling evidence that the trophic cascade effect is a real and widespread ecological phenomenon, rather than a coincidence.
Understanding Population Controls: Top-Down vs. Bottom-Up
- Elk populations, and populations in general, are subject to two primary types of controls:
- Top-Down Control: This refers to the influence of predators from "above" in the food chain. In the Yellowstone example, wolf predation (and the fear of it) controls elk behavior and numbers.
- Bottom-Up Control: This refers to the influence of resource availability from "below" in the food chain. For elk, this means the amount of available plant material (food) dictates population limits.
- Ecological thinking acknowledges that populations are influenced by a dynamic interplay of both top-down (predation) and bottom-up (food availability) trophic factors.
- Elk populations, and populations in general, are subject to two primary types of controls: