Countries including Iceland, Nicaragua, Senegal and Norway supported the proposal, claiming that the IWC would be "dysfunctional" if it did not approve it.

Opponents -  including Australia, Argentina and the IWC's 24 European Union member states -  argued that many whale populations are still vulnerable and that whaling is increasingly seen as unacceptable. They said that "disagreement does not mean dysfunction".

Australia was the first country to argue against the proposal. Australia's commissioner to the IWC Nick Gales described commercial whaling as "a business proposition against which many parties hold legitimate environmental and welfare concerns".

After the vote, Masaaki Taniai, Japan's state minister of agriculture, forestry and fisheries, said: "If scientific evidence and diversity are not respected, if commercial whaling based on science is completely denied, and if there is no possibility for the different positions and views to co-exist with mutual understanding and respect, then Japan will be pressed to undertake a fundamental reassessment of its position as a member of the IWC."

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Patrick Ramage, director of marine conservation at the International Fund for Animal Welfare, noted that Japan had frequently threatened to pull out of the body.

The measure's "adoption would have been a big step backwards for the IWC, returning us to the bygone days of open commercial whaling instead of becoming a modern conservation body", he said.Sea Shepherd said.

"The real way forward for whales is conservation and responsible whale watching, not cruel and unnecessary whale killing."

Alexia Wellbelove, senior program manager at animal charity Humane Society International in Australia said: "The IWC has a large, expanding and impressive marine conservation agenda, bringing countries together to find ways to protect whales from problems like bycatch and pollution.

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"Against that important and urgent backdrop, Japan's pro-whaling ambitions look regressive and vastly out of step with enlightened global efforts to act as stewards for the world's largest mammals.

"It's clear from exchanges this week that those countries here fighting for the protection of whales are not prepared to have the IWC's progressive conservation agenda held hostage to Japan's unreasonable whaling demands."

HSI president Kitty Block said: "It is an immense relief that the IWC's moral compass has led it to reject Japan's reckless and retrograde attempt to bring back commercial whaling. What Japan tried to do here was to bend and break the rules of the IWC to lift an internationally-agreed ban on killing whales for profit.

"It deserved to fail; the world has moved on from commercial whaling, and so must Japan. We hope that the IWC can now get on with the business of protecting these ocean leviathans from the myriad other threats they face."

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Sea Shepherd says that more than 32,000 whales have been killed by Japan, Iceland, Norway and Russia since the moratorium.

Sea Shepherd’s founder Captain Paul Watson, who has spent many years opposing Japan’s "scientific research" programs, said on Saturday: "Following yesterday’s most welcome Florianopolis Declaration, this defeat of the Japanese proposal has made the 67th meeting of the International Whaling Commission an awesome historical event for the world’s whales.

"The Florianopolis Declaration states that the purpose of the IWC is the conservation of whales and that the commercial killing of whales is to no longer be up for further discussion," he said.

On Wednesday the IWC adopted a number of measures protecting whales, including a resolution recognising their crucial role in ecosystems.

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While, in the past, the IWC has looked at whales simply as resources, the mammals are now seen as "ecosystem engineers", moving deep sea nutrients into the sunlight where their faecal plumes fertilise the water and help the productivity of ecosystems.

The resolution was passed in spite of opposition from Japan.

Two further resolutions - mitigating the threats among cetaceans of both noise pollution and getting entangled in ghost gear - were adopted by consensus on Wednesday.

In addition, the IWC's new bycatch initiative, which addresses a worldwide threat to whales and dolphins that are accidentally caught and drowned in fishing nets, was endorsed without a vote being necessary.

The day before, a proposal to create a whale sanctuary in the South Atlantic - backed among others by Australia, New Zealand and the hosts - did not receive enough support to pass.

"Whales use sound to help them navigate the vast oceans, but human-caused noise pollution can interfere with their ability to sense their environment and communicate with each other over vast distances," Ms Wellbelove said.

"This acoustic fog can have serious detrimental impacts on cetaceans, while other forms of noise pollution from industrial or military activity can even cause physical harm.

"Seeing these cetacean conservation measures pass at the IWC is a bittersweet experience. It is thrilling to see the IWC play such a globally important role in protecting cetaceans in their marine environment and recognising the vital and unique role these ocean giants play in keeping our seas healthy.

"Yet all the while there are moves at the IWC by Japan and others to kill these very same whales for profit."

With AP