Species → Ochre sea star
Ochre sea star
Pisaster ochraceus
What if it disappeared?
Impact MajorThe number of species on the intertidal shore drops from 15 to 8: without a predator, the mussel Mytilus californianus takes over the available space and crowds out the species that competed with it for rock attachment sites.
DocumentedDetail
Paine, R.T. (1966). Food Web Complexity and Species Diversity. The American Naturalist, 100(910), 65–75.
By experimentally removing the ochre sea star from plots of rocky intertidal shore and comparing them to control plots, Paine showed that the mussel, the sea star's main prey, comes to monopolize the available space within three years, crowding out seven other rock-attached species. This is the founding experiment behind the keystone species concept.
The shore becomes a mussel monoculture — a single species occupies nearly all the space once shared among several.
DocumentedDetail
Lafferty, K.D. et Suchanek, T.H. (2016). Revisiting Paine's 1966 Sea Star Removal Experiment, the Most-Cited Empirical Article in the American Naturalist. The American Naturalist, 188(4), 365–378.
This revisit of Paine's experimental site, several decades later, confirms that over the long term, with the sea star durably absent, the mussel eventually forms an almost continuous mat — even though, when counting the species living within the mussel bed itself (worms, small crustaceans), the site's total diversity otherwise remains high.
The real-world decline of the sea star across large portions of the Pacific coast has not been the subject of published monitoring as complete as Paine's controlled experiment.
InsufficientDetail
Wasting disease caused sea star density and biomass to drop by 59 to 90% depending on the site between 2013 and 2014, but recovery has been very uneven across regions (better in Oregon and northern California than in Alaska and Washington). No one has confirmed whether the mussel takeover Paine observed at small scale has occurred at the regional scale of this real-world epidemic.
Documentation
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Sea Star Wasting Disease in the Keystone Predator Pisaster ochraceus in Oregon
Accessed August 10, 2026
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Paine (1966) — Food Web Complexity and Species Diversity, The American Naturalist
Accessed August 7, 2026
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Lafferty & Suchanek (2016) — Revisiting Paine's 1966 Sea Star Removal Experiment, The American Naturalist
Accessed August 7, 2026
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Miner et al. (2018) — Large-scale impacts of sea star wasting disease (SSWD) on intertidal sea stars, PLOS ONE
Accessed August 7, 2026
Last verified: August 14, 2026