Flagship Scottish daily The Scotsman now has more readers online than in print, according to figures released today.

The National Readership Survey has for the first time published data combining print and online audiences for national newspapers, along with a handful of regional titles.

The Scotsman had an average of 513,000 online readers compared to 459,000 print readers a month, giving the Johnston Press title a combined total of 940,000 and making it the only regional newspaper with more digital readers than print.

The NRS figures, which are calculated from surveys with individuals, relate to the year from April 2011 to March 2012 with the online data coming from UKOM/Nielsen for April 2012.

Glasgow-based The Herald had a combined readership of 697,000 a month, with 225,000 of these from its website, while its Newsquest sister title the Glasgow Evening Times had combined figures of 513,000, with 65,000 being digital readers.

The Yorkshire Post had a combined total of 907,000 readers, with 241,000 of these coming from its website, while the London Evening Standard had 4.5m print readers and 531,000 onlines ones.

Dundee-based The Courier and Advertiser had 297,000 print readers and 96,000 online ones, giving it a combined total of 387,000.

The NRS figures are calculated differently from the ABC sales figures and are based on a survey of 35,000 individuals.