Based on detailed documentation of the state of mammals, birds, fish and butterflies on the island, it found that at least 881 of 3,196 recorded species, or 28 per cent, had vanished forever.

However, taking into account the probable number of animals here before detailed records were made in the late 1800s, it predicted that this figure was actually higher - about half of Singapore's animals.

The study, done by researchers from the National University of Singapore (NUS) and Australia's Northern Territory University, is being published today in the prestigious science journal Nature, an international weekly which is read by researchers and scientists.

It is one of a handful of such studies based on data gathered over such a long period.

By looking at the extensive habitat loss in Singapore and the current rate at which forests in the region are being felled, it predicted the loss of up to 42 per cent of animal populations in South-east Asia by the end of this century.

At least half will be species unique to this region.

'Clearly, large-scale conservation efforts need to be implemented if these rates of extinction are to be abated,' it said.

It added that in Singapore, since the British first established a presence here in the early 1800s, more than 95 per cent of the estimated 540 sq km of original vegetation has been cleared.

Now, nature reserves, which make up only 0.25 per cent of Singapore's land area, are home to more than half of the native animals here.

Rapid, large-scale habitat destruction for agriculture and urban development were the main culprits behind the mass extinctions.

But over-hunting and fishing, and the heavy shelling of nature reserves during World War II, also played a role.

Some of the animals most threatened with extinction are two mammals found only in Singapore - the cream- coloured giant squirrel and banded leaf monkey, said Associate Professor Peter Ng of the NUS department of biological sciences, one of the authors of the study.

Sadly, they are almost certain to become extinct because their populations have shrunk to levels too small for them to be sustainable in the long run.

'These are effectively the living dead,' said Prof Ng, who is also director of the Raffles Museum of Biodiversity Research.

The prospects for Singapore's surviving animals look bleak, said the paper, pointing out that the 77 per cent of the species are under threat, and most of them reside in our nature reserves.

But all is not lost.

Said NUS Associate Professor Navjot S. Sodhi, a bird expert and co-author of the paper: 'Hopefully, South-east Asia can take some lessons from what has happened in Singapore, and do something to better protect its animals.'

He added that without Singapore's nature reserves, the results here would have been even bleaker.

Nee Soon Reserve, for example, is home to one-quarter of the remaining freshwater crab and fish here.

Said Prof Ng: 'It's imperative to hang on to all our protected areas.

'We can't change history, but we can try to hang on to what we have now.'