Aberdeen and Dundee recorded the largest annual falls of 0.7 per cent each, while Midlothian and East Lothian saw the biggest population increases, of 1.6 per cent each.

The estimates for the year to June 2021 – the first for a full year of the pandemic – also show the populations of Edinburgh and Glasgow fell, but areas like Aberdeenshire and Argyll and Bute recorded their first increase for years.

Factors involved could include people moving to find more space to work from home, and students moving back in with their parents during the lockdowns or foreign students returning home.

Dundee saw the equal largest population fall among Scottish cities. Picture: John Devlin

Scotland’s overall population is estimated to have increased 0.25 per cent or by 13,900 people to 5,479,900 but the growth rate was slower than the previous five years and is expected to start to fall by the end of the decade.

The rise continues to be principally fuelled by migration, following a 20-year trend, with more people moving to Scotland than those leaving.

By contrast, there have been more deaths than births for seven years, with the gap between them now the largest on record.

Esther Roughsedge, head of population and migration statistics at NRS, said: “As well as people moving long-term out of cities and into the surrounding areas, there may have been students who have moved back to their parents’ addresses temporarily during the pandemic.

“Another factor could be people who had previously moved updating their address with a GP to make sure they received their Covid vaccination letters.

“However, our most recent projections looking ahead to 2045, published in January, show that if current trends in births, deaths and migration continue, Scotland’s population will start to fall by the end of this decade.”

Dr Sarah Christison, a post-doctoral research fellow in the school of geography and sustainable development at the University of St Andrews, said the city-rural population changes “could be due to an increase in remote working or working from home where people have greater freedom to further from their workplace.

"Lower numbers of international students present in Scotland’s cities while online learning was in place may also have an effect.

"We can see in Dundee, which has one of the largest student populations in Scotland, also experienced the largest decline in population.

“Changes in the costs of living may have also influenced people’s decision to move from cities to more rural areas.

"Combined with greater flexibility as a result of remote working, more affordable housing and living costs may have drawn people from large cities to neighbouring, more rural areas.”

However, she added: “More data would be needed before we could say if these changes are a short-term shift as an immediate result of the pandemic or whether there is likely to be longer-term trend towards rural population growth and urban population decline.”

Robert Wright, a consultant and former professor of economics at the University of Strathclyde, said: “Just blame Brexit. Brexit and Covid have reduced immigrants. Since most immigrants end up in cities, it is not surprising that city growth has slowed.”

Professor Michael Anderson of the University of Edinburgh, and author of Scotland's Populations from the 1850s to Today, said: "The cities suggest to me a big student-loss effect, with many going home to rural areas whence they would normally have left.

"Also, second home owners moving to their second homes such as in Argyll and Bute, Highland, and Perth and Kinross seems likely and, critically, re-registering there with their GPs, which is the source of the figures.

"But the key point is that this is a disrupted year, with almost certainly less consistent statistics, and unlikely to be a good guide for the future.”

A Glasgow City Council spokesperson said: "Glasgow’s population fell by less than one in a thousand – the first year the population did not rise in the city since the modern low of 578,000 in 2001, and the lowest decrease of all the Scottish cities.

“As working, socialising and studying arrangements move closer to ‘normal’ post-pandemic, it could reasonably be expected that the two-decade trend will recommence.”

An Aberdeen City Council spokesperson said: “Aberdeen has faced some very challenging times in recent years with the downturn in the oil and gas industry allied to the unprecedented challenge of the Covid pandemic.

"These two factors have undoubtedly had a negative impact on the local economy and in turn contributed to a dip in the population.

"We are nothing if not resilient however, and we are actively undertaking work to encourage city living with new high-quality council housing developments, schools, community hubs and other projects designed to make the city an even more desirable place to live.”

A spokesperson for Dundee City Council said: “Along with our partners, we are continuing to work to maintain a strong and sustainable city economy that will provide jobs for the people of Dundee, retain more graduates and make the city a magnet for new talent.”

Argyll and Bute Council said: “Our location and lifestyle can be key attractions for people.

"We are close to the Central Belt of Scotland and at the same time provide a popular outdoors lifestyle.

"With working from home increasing, moving here for the work/life balance our location gives is an option for many.”

An Aberdeenshire Council spokesperson said: “There is certainly a significant trend of people moving from cities to rural areas, taking advantage of reasonably-priced housing, more flexible and remote work opportunities and the greater importance placed on quality of life.

“That is something we enjoy here in Aberdeenshire and when you consider the wide range of excellent job opportunities, strong transport connections, excellent schools and our outstanding natural environment it is perhaps not surprising that we are seeing this recent increase in the population.”

Covid reverses long-term population trend as rural areas benefit at expense of Scotland’s cities

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