Thousands of Leave voters deserted the SNP in June’s general election thanks to Nicola Sturgeon’s pro-EU stance and attempts to force a second independence referendum, according to an academic study into the party’s mauling.

Research conducted by the University of Manchester found that four out of ten Scots who backed independence in 2014 and leaving the EU in last year’s referendum switched support from the SNP to another party.

It said that many people who backed the SNP in the 2015 election, when the party won 56 out of 59 Scottish seats, were “driven away by the party’s strong pro-Remain stance.”

Ruth Davidson’s Conservatives and Kezia Dugdale’s Scottish Labour attracted similar proportions of disillusioned former SNP supporters who backed Leave, the report found.

But the Tories also hoovered up Unionist voters who had backed Labour in the 2015 election, especially among those who supported Leave in the EU referendum.

Academics said that the ability of the Scottish Conservatives to capture Unionists who voted both Remain and Leave could be partly attributed to their “strong campaign.”

However, it also said their success reflects the fact that voters identified more closely with how they cast their ballots in the independence referendum than the EU referendum.