Martin Friend, 24, from Orpington, Kent, Steven Roberts, 27, from Edinburgh, Mark Lindgren, 23, from St Albans, Hertfordshire, and Joanne Cotton, 28, from Nazeing, Essex, were beaten to death by Hutu rebels on the first day of their holiday.

Mark Lindgren, one of those beaten to death

Four other tourists and four guides were also killed in the raid on a tourist camp at Bwindi national park on 1 March. Another six tourists survived the attack.

The inquest in Crawley, West Sussex, heard the four dead had suffered severe head injuries and had each been hit with a heavy blunt instrument up to five times.

Pathologist Dr Iain West told the hearing they would have been knocked unconscious by the first blow. He said they had not been hacked, mutilated or subject to any sexual attack.

Rebels were men and women

A Metropolitan police officer sent to Uganda after the massacre told the inquest the rebels had stormed the tourist camp at about 0630 local time, while most of the tourists were sleeping.

Vehicles were destroyed so the alarm could not be raised

Detective Superintendent Ken Woodward said the group attacking the camp were men and women, all armed with rifles, handguns, machetes and wooden clubs.

They were believed to be from the ragtag Interahamwe militia, whose members have been based in the neighbouring Democratic Republic of Congo since their defeat by the Rwandan Patriotic Front in 1994.

Det Supt Woodward said the rebels rounded up the visitors, looted property, clothing, cameras and jewellery, and took rifles and ammunition from park rangers.

The attack was launched from the neighbouring DR Congo

One local ranger was killed as he tried to defend the camp, and several rangers and tourists escaped the attack by hiding or fleeing.

Marched in silence, barefoot

Mr Woodward described how the tourists were separated into groups of different nationalities.

The rebels then selected a group of 16, all Westerners, and one local ranger to be taken up a trail towards the Congo border, five miles (seven kilometres) away.

Mark Avis, who survived, and wife Rhonda, who died

Americans Rob Haubner, 48, and his wife Susan Miller, 42, were killed about 1.5km from the camp when they were killed.

Ms Cotton, New Zealanders Michelle Strathern, 26, and Rhonda Avis, 27, whose husband Mark survived, were also killed on the trail.

Intention 'was only to loot'

Mr Woodward said that he believed that the intial intention of the Interahamwe had been merely to loot the village.

He said: "We will never know what sparked the original deaths and subsequent deaths of the others. It may have been that they found it difficult to travel in bare feet and lagged behind.

"Or maybe the intention was too kill them for the maximum publicity around the world."

Mr Woodward said three handwritten messages had been left by the rebels - one was given to the released hostages and the others were left near the bodies or on the trail.

He did not read the messages to the court but said they showed "a hatred for Anglo-Saxons and Americans."

Decoy attack

It has been speculated that the killings were in revenge against the US and UK governments for allegedly supporting the rebels' Tutsi rivals in the Rwandan government.

Mr Woodward told the inquest that there had been no history of terrorist activity at Bwindi before the attack.

Mr Woodward added: "It is unlikely that now anyone from that Interahamwe group will be caught and prosecuted.

Deaths a 'complete disaster'

Recording a verdict of unlawful killing, West Sussex coroner Roger Stone said the deaths were "a complete disaster" for the families, many of whom were at the inquest.

"It will take many years, if indeed, ever, to come to terms with the deaths," he said.

Uganda's President Yoweri Museveni pledged to take revenge on the rebels in the wake of the attack.

There was a massive manhunt by the Ugandan army and reported gun battles, in which large numbers of the rebels are understood to have been killed.

A special committee with responsibility for tourist safety was set up to work on further measures to prevent such an incident happening in Uganda again.

The Bwindi and Mgahinga parks were reopened in April after being closed for a month of mourning.