Syria's conflict has killed tens of thousands and displaced millions, but it has also seen the country's irreplaceable heritage dug up and sold, damaged by war and wilfully destroyed.
From pre-historic settlements and ancient markets, to Greco-Roman ruins and Crusader castles, Syria is a repository of treasures from different ages and cultures.
But that rich history has also made the country's war particularly devastating from the perspective of archaeologists and heritage specialists.
"There are more than 900 monuments and archaeological sites that have been affected, destroyed or wiped out," said Cheikhmous Ali of the Association for Protection of Syrian Archaeology.
"And that is something of concern not just for Syria, because this is international heritage."
On Monday, satellite images confirmed the Temple of Bel in the ruins of ancient Palmyra had been destroyed by the Islamic State group just a week after the jihadists levelled the smaller Baal Shamin temple.
The loss is particularly painful, as Palmyra is one of Syria's six UNESCO World Heritage-listed sites, described by the United Nations as "of incalculable value to our shared global heritage".
The recent destruction in Palmyra, which experts said was likely to continue, has been some of the most wanton in the war in Syria.
The Islamic State group's harsh philosophy condemns pre-Islamic religious sites and considers statues and grave markers to be idolatrous.
But the group has also been accused of targeting heritage to loot items for the black market and to gain publicity.
- 'They have killed Palmyra' -
IS has been quick to destroy sites and artefacts that fall outside its interpretation of religion and history.
Muslim sites including the Sufi Uwais al-Qarni Mosque and a shrine to a companion of the Prophet Mohammed have been targeted in the group's stronghold of Raqa, along with Christian sites like the Mar Elian monastery in the central province of Homs.
Before IS arrived in Palmyra in late May, antiquities officials were able to evacuate its museum, removing many priceless pieces.
But Syria's antiquities chief Maamoun Abdulkarim said several remaining sites and pieces that could not be moved were now under threat.
"There are still dozens of the greatest graves, the amphitheatre and the Temple of Nabu," he told AFP.
"They have killed Palmyra. Now, they will terrorise it," he said.
Maurice Sartre, a French archaeologist and expert on Palmyra, was furious about the destruction of the Temple of Bel and accused the West of inaction.
"My personal feeling is that of huge anger against Daesh," he said using another acronym for IS.
"I didn't expect anything else from them... but also against all European and American political leaders who remain almost motionless."
The grand amphitheatre has already been used for a gruesome massacre in which child members of IS killed 25 Syrian soldiers, execution-style, in front of residents.
While the devastation at Palmyra is perhaps the most brazen in the conflict, experts have long warned of the piecemeal destruction at other heritage sites.
In December 2014, the UN said nearly 300 important sites had been destroyed, damaged or looted since the conflict began in March 2011.
Using satellite imaging systems, the UN said 24 sites had been completely destroyed.
Another 104 were severely damaged, with 85 moderately damaged and 77 possibly damaged, the survey found.
Among the areas studied were Syria's six UNESCO World Heritage-listed sites: the ancient cities in Aleppo, Damascus and Bosra, the Dead Cities of northern Syria, the Crac des Chevaliers castle and the ruins of Palmyra.
- 'Systematic destruction' -
Much of the damage to Syria's heritage sites has been caused by heavy fighting.
The northern city of Aleppo, where settlement dates back 7,000 years, has been ravaged by three years of clashes between rebels and the government.
Its ancient market has been badly damaged by fighting and fire, and the minaret of the 11th century Great Mosque of Aleppo has been reduced to rubble.
Similarly, the Crac des Chevaliers Crusader castle suffered serious damage to its facade and roof before government forces recaptured it from rebels.
Elsewhere, damage has been caused by looters, who have taken advantage of the lawlessness of war to ramp up their activities to devastating levels.
Even before IS's arrival, looters were digging up items at Palmyra to be smuggled for sale abroad.
The UN last year said the Dura-Europos site in eastern Deir Ezzor province, once known as the "Pompeii of the desert" had been rendered "unrecognisable" by looting.
Ali, whose organisation documents the destruction to Syria's heritage, said the damage was just part of the unravelling of his country.
"Syria is being destroyed in every field, whether infrastructure, humanitarian affairs, relations between communities or heritage," he said.
"It is a systematic destruction that has gone on for four years