WASHINGTON — The dwindling march of the penguins is signaling that the world’s oceans are in trouble, scientists now say.

Penguins may be the tuxedo-clad version of a canary in the coal mine, with generally ailing populations from a combination of global warming, ocean oil pollution, depleted fisheries, and tourism and development, according to a new scientific review paper.

University of Washington biologist P. Dee Boersma detailed specific problems around the world with remote penguin populations, linking their decline to the overall health of southern oceans.

Scientists figure there are between 16 to 19 species of penguins. About a dozen are in some form of trouble, Boersma wrote.

The largest Patagonian penguin colony in the world is at Punta Tumbo, Argentina, but the number of breeding pairs there dropped in half from about 400,000 in the late 1960s to about 200,000 in October 2006, Boersma reported. Over a century, African penguins have decreased from 1.5 million breeding pairs to 63,000.

The decline overall isn’t caused by one factor, but several.

For the ice-loving Adelie penguins, for example, global warming in the western Antarctica peninsula is a problem, making it harder for them to find food, said Phil Trathan, a top penguin scientist.

For penguins that live on the Galapagos island, El Nino weather patterns are a problem because the warmer water makes penguins travel farther for food, at times abandoning their chicks, Boersma said.