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The most startling change in combined readership was shown by The Sun, thanks to it dropping its paywall completely in November 2015, according to the latest readership estimates from Pamco.

The News UK tabloid's combined readership jumped by 91% to 26,196,000, putting it ahead of the Daily Mirror and not far off the MailOnline, the Daily Mail's companion website.

Last month ±±¾©Èü³µpk10 revealed The Sun had overtaken The Mirror, based on figures from ComScore.

But The Sun’s PC growth of 56% to 2,780,000 and eight-fold mobile growth to 18,956,000 may have come at the expense of its print readership, which dropped 10% to 10,123,000.

Pamco's print and digital estimates allow a comparison of publishers' 2015 and 2016 monthly average readership figures. 

There was a twisted echo of that trend at the overall newsbrand market level. Print readership dropped by 3% and mobile grew by 4%, but the biggest change was in PC readership, which dropped by 8%.

This meant only 1% growth for newsbrand readership across all platforms, which left only Daily Mail owner DMGT unaffected by the draw of The Sun, and a lesser extent, The Times.

Speculation about the Daily Express and Daily Star being sold to Trinity Mirror will not be lessened by their respective combined readership drops of 7% and 9%.

Both titles experienced double-digit percentage declines in mobile readership, in contrast to the Daily Mirror’s 3% rise.

However, the Mirror lost readers on PC and in print, leaving its combined readership down by 1%.

At the other end of the market the momentum is with The Times, which slightly softened its hard paywall in July 2016 to allow non-subscribers to read two stories per week in return for registering.

This helped boost its combined readership by 16% to 5,790,000.

However, those extra 780,000 Times readers are hard to square with the ten million readers lost by three rivals: The Guardian, The Independent and The Telegraph.

The Telegraph lost nearly five million, 19% of its combined readership, and the blame was down to its PC and mobile platforms.

In November last year, too late to have much effect on the figures, it opted to switch its metered paywall for a hybrid free and subscription content approach.

The Guardian also saw its stable print readership undermined by its digital performance despite its free access website, and lost 8 per cent of its combined readership.

The Independent also lost digital readers even though it dropped its print edition in March 2016. Readership fell on PC by 17% to 4,376,000 and on mobile by 2% to 14,405,000.

The Pamco figures also cover a number of magazines, with the medium experiencing a similar move to mobile but a slight decline in combined readership to 37,691,000.

A News UK spokeswoman said: "We're immensely proud of the total brand reach of The Sun, The Sun on Sunday and thesun.co.uk. The digital team under Keith Poole have grown The Sun website from a standing start to a scaled audience in the UK and there aren't many sites that have the skill set and brand recognition to be able to do that.

"Nearly half of all of our readers choose to read the print product on a monthly basis, which means that The Sun remains the most widely-read newspaper in the UK."