London was by far the largest urban agglomeration in the United Kingdom in 2025, with an estimated population of *** million people, more than three times as large as Manchester, the UK’s second-biggest urban agglomeration. The agglomerations of Birmingham and Leeds / Bradford had the third and fourth-largest populations, respectively, while the biggest city in Scotland, Glasgow, was the fifth largest. Largest cities in Europe Two cities in Europe had larger urban areas than London, with Istanbul having a population of around **** million and the Russian capital Moscow having a population of over **** million. The city of Paris, located just over 200 miles away from London, was the second-largest city in Europe, with a population of more than **** million people. Paris was followed by London in terms of population size, and then by the Spanish cities of Madrid and Barcelona, at *** million and *** million people, respectively. The Italian capital, Rome, was the next largest city at *** million, followed by Berlin at *** million. London’s population growth Throughout the 1980s, the population of London fluctuated from a high of **** million people in 1981 to a low of **** million inhabitants in 1988. During the 1990s, the population of London increased once again, growing from ****million at the start of the decade to **** million by 1999. London's population has continued to grow since the turn of the century, and despite declining between 2019 and 2021, it reached *** million people in 2023 and is forecast to reach almost *** million by 2047.

This statistic shows the ten largest cities in the United Kingdom in 2021. In 2021, around 8.78 million people lived in London, making it the largest city in the United Kingdom.

National and subnational mid-year population estimates for the UK and its constituent countries by administrative area, age and sex (including components of population change, median age and population density).

Cambridge was the fastest growing city in the United Kingdom between 2013 and 2023, with its population increasing by 17.3 percent. Exeter, Milton Keynes, and Peterborough also grew quite fast, with their populations increasing by 15.2 percent, 14.9 percent, and 14 percent, respectively. Largest UK urban areas When looking at cities defined by their urban agglomerations, as of 2023, London had approximately 9.65 million people living there, far larger than any other city in the United Kingdom. The urban agglomeration around the city of Birmingham had a population of approximately 2.67 million, while the urban areas around Manchester and Leeds had populations of 2.79 and 1.92 million respectively. London not only dominated other UK cities in terms of its population, but in its importance to the UK economy. In 2023, the gross domestic product of Greater London was approximately 569 billion British pounds, compared with 101 billion for Greater Manchester, and 85 billion in the West Midlands Metropolitan Area centered around Birmingham. UK population growth In 2023, the overall population of the United Kingdom was estimated to have reached approximately 68.3 million, compared with around 58.9 million in 2000. Since 1970, 2023 was also the year with the highest population growth rate, growing by 0.98 percent, and was at its lowest in 1982 when it shrank by 0.12 percent. Although the UK's birth rate has declined considerably in recent years, immigration to the UK has been high enough to drive population growth in the UK, which has had a positive net migration rate since 1994.

In 2025, Moscow was the largest city in Europe with an estimated urban agglomeration of 12.74 million people. The French capital, Paris, was the second largest city in 2025 at 11.35 million, followed by the capitals of the United Kingdom and Spain, with London at 9.84 million and Madrid at 6.81 million people. Istanbul, which would otherwise be the largest city in Europe in 2025, is excluded as it is only partially in Europe, with a sizeable part of its population living in Asia. Europe’s population is almost 750 million Since 1950, the population of Europe has increased by approximately 200 million people, increasing from 550 million to 750 million in these seventy years. Before the turn of the millennium, Europe was the second-most populated continent, before it was overtaken by Africa, which saw its population increase from 228 million in 1950 to 817 million by 2000. Asia has consistently had the largest population of the world’s continents and was estimated to have a population of 4.6 billion. Europe’s largest countries Including its territory in Asia, Russia is by far the largest country in the world, with a territory of around 17 million square kilometers, almost double that of the next largest country, Canada. Within Europe, Russia also has the continent's largest population at 145 million, followed by Germany at 83 million and the United Kingdom at almost 68 million. By contrast, Europe is also home to various micro-states such as San Marino, which has a population of just 30 thousand.

All cities with a population > 1000 or seats of adm div (ca 80.000)Sources and ContributionsSources : GeoNames is aggregating over hundred different data sources. Ambassadors : GeoNames Ambassadors help in many countries. Wiki : A wiki allows to view the data and quickly fix error and add missing places. Donations and Sponsoring : Costs for running GeoNames are covered by donations and sponsoring.Enrichment:add country name

As of 2023, the population density in London was by far the highest number of people per square km in the UK, at *****. Of the other regions and countries which constitute the United Kingdom, North West England was the next most densely populated area at *** people per square kilometer. Scotland, by contrast, is the most sparsely populated country or region in the United Kingdom, with only ** people per square kilometer. Countries, regions, and cities According to the official mid-year population estimate, the population of the United Kingdom was just almost **** million in 2022. Most of the population lived in England, where an estimated **** million people resided, followed by Scotland at **** million, Wales at **** million and finally Northern Ireland at just over *** million. Within England, the South East was the region with the highest population at almost **** million, followed by the London region at around *** million. In terms of urban areas, Greater London is the largest city in the United Kingdom, followed by Greater Manchester and Birmingham in the North West and West Midlands regions of England. London calling London's huge size in relation to other UK cities is also reflected by its economic performance. In 2021, London's GDP was approximately *** billion British pounds, almost a quarter of UK GDP overall. In terms of GDP per capita, Londoners had a GDP per head of ****** pounds, compared with an average of ****** for the country as a whole. Productivity, expressed as by output per hour worked, was also far higher in London than the rest of the country. In 2021, London was around **** percent more productive than the rest of the country, with South East England the only other region where productivity was higher than the national average.

According to the 2021 Census, London was the most ethnically diverse region in England and Wales – 63.2% of residents identified with an ethnic minority group.

The population of the United Kingdom in 2023 was estimated to be approximately 68.3 million in 2023, with almost 9.48 million people living in South East England. London had the next highest population, at over 8.9 million people, followed by the North West England at 7.6 million. With the UK's population generally concentrated in England, most English regions have larger populations than the constituent countries of Scotland, Wales, and Northern Ireland, which had populations of 5.5 million, 3.16 million, and 1.92 million respectively. English counties and cities The United Kingdom is a patchwork of various regional units, within England the largest of these are the regions shown here, which show how London, along with the rest of South East England had around 18 million people living there in this year. The next significant regional units in England are the 47 metropolitan and ceremonial counties. After London, the metropolitan counties of the West Midlands, Greater Manchester, and West Yorkshire were the biggest of these counties, due to covering the large urban areas of Birmingham, Manchester, and Leeds respectively. Regional divisions in Scotland, Wales and Northern Ireland The smaller countries that comprise the United Kingdom each have different local subdivisions. Within Scotland these are called council areas whereas in Wales the main regional units are called unitary authorities. Scotland's largest Council Area by population is that of Glasgow City at over 622,000, while in Wales, it was the Cardiff Unitary Authority at around 372,000. Northern Ireland, on the other hand, has eleven local government districts, the largest of which is Belfast with a population of around 348,000.

According to the 2021 Census, there were 30.4 million (51.0%) women and girls and 29.2 million (49.0%) men and boys in England and Wales.

Mid-year population estimates for Glasgow available via NOMIS API link. This data is provided via the API from NOMIS Licence: None Mid-Year Population Estimates for the last 10 years.json - https://dataservices.open.glasgow.gov.uk/Download/Organisation/41792b4a-74db-407b-a8b5-cc6695cb7151/Dataset/ed007e60-e492-459b-acbb-ef23a29d24cc/File/bc3f0778-ddfb-403e-97b3-9fb032d0bc5a/Version/0e37e605-7081-4cce-9164-e80012dfb11e

This report was released in September 2010. However, recent demographic data is available on the datastore - you may find other datasets on the Datastore useful such as: GLA Population Projections, National Insurance Number Registrations of Overseas Nationals, Births by Birthplace of Mother, Births and Fertility Rates, Office for National Statistics (ONS) Population Estimates

FOCUSONLONDON2010:POPULATIONANDMIGRATION

London is the United Kingdom’s only city region. Its population of 7.75 million is 12.5 per cent of the UK population living on just 0.6 per cent of the land area. London’s average population density is over 4,900 persons per square kilometre, this is ten times that of the second most densely populated region.

Between 2001 and 2009 London’s population grew by over 430 thousand, more than any other region, accounting for over 16 per cent of the UK increase.

This report discusses in detail the population of London including Population Age Structure, Fertility and Mortality, Internal Migration, International Migration, Population Turnover and Churn, and Demographic Projections.

Population and Migration report is the first release of the Focus on London 2010-12 series. Reports on themes such as Income, Poverty, Labour Market, Skills, Health, and Housing are also available.

REPORT:

Read the full report in PDF format.

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PRESENTATION:

To access an interactive presentation about population changes in London click the link to see it on Prezi.com

DATA:

To access a spreadsheet with all the data from the Population and Migration report click on the image below.

Report data

MAP:

To enter an interactive map showing a number of indicators discussed in the Population and Migration report click on the image below.

Interactive Maps

FACTS:

● Top five boroughs for babies born per 10,000 population in 2008-09:

1. Newham – 244.4
2. Barking and Dagenham – 209.3
3. Hackney – 205.7
4. Waltham Forest – 202.7
5. Greenwich – 196.2

-32. Havering – 116.8

-33. City of London – 47.0

● In 2009, Barnet overtook Croydon as the most populous London borough. Prior to this Croydon had been the largest since 1966

● Population per hectare of land used for Domestic building and gardens is highest in Tower Hamlets

● In 2008-09, natural change (births minus deaths) led to 78,000 more Londoners compared with only 8,000 due to migration. read more about this or click play on the chart below to reveal how regional components of populations change have altered over time.

This statistic shows the ten cities with the highest number of businesses per 10,000 population in the United Kingdom (UK) in 2018. London had the highest number of businesses per 10,000 population in this year at ***, followed by Milton Keynes which had ***.

By 1800, London had grown to be the largest city in Western Europe with just under one million inhabitants. Paris was now the second largest city, with over half a million people, and Naples was the third largest city with 450 thousand people. The only other cities with over two hundred thousand inhabitants at this time were Vienna, Amsterdam and Dublin. Another noticeable development is the inclusion of many more northern cities from a wider variety of countries. The dominance of cities from France and Mediterranean countries was no longer the case, and the dispersal of European populations in 1800 was much closer to how it is today, more than two centuries later.

The empirical dataset is derived from a survey carried out on 25 estates in 14 cities in nine different European countries: France (Lyon), Germany (Berlin), Hungary (Budapest and Nyiregyha´za), Italy (Milan), the Netherlands (Amsterdam and Utrecht), Poland (Warsaw), Slovenia (Ljubljana and Koper), Spain (Barcelona and Madrid), and Sweden (Jo¨nko¨ping and Stockholm). The survey was part of the EU RESTATE project (Musterd & Van Kempen, 2005). A similar survey was constructed for all 25 estates.

The survey was carried out between February and June 2004. In each case, a random sample was drawn, usually from the whole estate. For some estates, address lists were used as the basis for the sample; in other cases, the researchers first had to take a complete inventory of addresses themselves (for some deviations from this general trend and for an overview of response rates, see Musterd & Van Kempen, 2005). In most cities, survey teams were hired to carry out the survey. They worked under the supervision of the RESTATE partners. Briefings were organised to instruct the survey teams. In some cases (for example, in Amsterdam and Utrecht), interviewers were recruited from specific ethnic groups in order to increase the response rate among, for example, the Turkish and Moroccan residents on the estates. In other cases, family members translated questions during a face-to-face interview. The interviewers with an immigrant background were hired in those estates where this made sense. In some estates it was not necessary to do this because the number of immigrants was (close to) zero (as in most cases in CE Europe).

The questionnaire could be completed by the respondents themselves, but also by the interviewers in a face-to-face interview.

Data and Representativeness

The data file contains 4756 respondents. Nearly all respondents indicated their satisfaction with the dwelling and the estate. Originally, the data file also contained cases from the UK.

However, UK respondents were excluded from the analyses because of doubts about the reliability of the answers to the ethnic minority questions. This left 25 estates in nine countries. In general, older people and original populations are somewhat over-represented, while younger people and immigrant populations are relatively under-represented, despite the fact that in estates with a large minority population surveyors were also employed from minority ethnic groups. For younger people, this discrepancy probably derives from the extent of their activities outside the home, making them more difficult to reach. The under-representation of the immigrant population is presumably related to language and cultural differences. For more detailed information on the representation of population in each case, reference is made to the reports of the researchers in the different countries which can be downloaded from the programme website. All country reports indicate that despite these over- and under-representations, the survey results are valuable for the analyses of their own individual situation.

This dataset is the result of a team effort lead by Professor Ronald van Kempen, Utrecht University with funding from the EU Fifth Framework.

This dataset represents ethnic group (19 tick-box level) by highest level qualification, for England and Wales combined. The data are also broken down by age and by sex.

The ethnic group that the person completing the census feels they belong to. This could be based on their culture, family background, identity, or physical appearance. Respondents could choose one out of 19 tick-box response categories, including write-in response options.

Total counts for some population groups may not match between published tables. This is to protect the confidentiality of individuals' data. Population counts have been rounded to the nearest 5 and any counts below 10 are suppressed, this is signified by a 'c' in the data tables.

"Asian Welsh" and "Black Welsh" ethnic groups were included on the census questionnaire in Wales only, these categories were new for 2021.

This dataset provides Census 2021 estimates that classify usual residents in England and Wales by ethnic group. The estimates are as at Census Day, 21 March 2021. This dataset shows population counts for usual residents aged 16+ Some people aged 16 years old will not have completed key stage 4 yet on census day, and so did not have the opportunity to record any qualifications on the census.

These estimates are not comparable to Department of Education figures on highest level of attainment because they include qualifications obtained outside England and Wales.

For quality information in general, please read more from here.

Ethnic Group (19 tick-box level)

These are the 19 ethnic group used in this dataset:

• Asian, Asian British or Asian Welsh
  • Bangladeshi
  • Chinese
  • Indian
  • Pakistani
  • Other Asian
• Black, Black British, Black Welsh, Caribbean or African
  • African
  • Caribbean
  • Other Black
• Mixed or Multiple ethnic groups
  • White and Asian
  • White and Black African
  • White and Black Caribbean
  • Other Mixed or Multiple ethnic groups
• White
  • English, Welsh, Scottish, Northern Irish or British
  • Gypsy or Irish Traveller
  • Irish
  • Roma
  • Other White
• Other ethnic group
  • Arab
  • Any other ethnic group

No qualifications

No qualifications

Level 1

Level 1 and entry level qualifications: 1 to 4 GCSEs grade A* to C , Any GCSEs at other grades, O levels or CSEs (any grades), 1 AS level, NVQ level 1, Foundation GNVQ, Basic or Essential Skills

Level 2

5 or more GCSEs (A* to C or 9 to 4), O levels (passes), CSEs (grade 1), School Certification, 1 A level, 2 to 3 AS levels, VCEs, Intermediate or Higher Diploma, Welsh Baccalaureate Intermediate Diploma, NVQ level 2, Intermediate GNVQ, City and Guilds Craft, BTEC First or General Diploma, RSA Diploma

Apprenticeship

Apprenticeship

Level 3

2 or more A levels or VCEs, 4 or more AS levels, Higher School Certificate, Progression or Advanced Diploma, Welsh Baccalaureate Advance Diploma, NVQ level 3; Advanced GNVQ, City and Guilds Advanced Craft, ONC, OND, BTEC National, RSA Advanced Diploma

Level 4 +

Degree (BA, BSc), higher degree (MA, PhD, PGCE), NVQ level 4 to 5, HNC, HND, RSA Higher Diploma, BTEC Higher level, professional qualifications (for example, teaching, nursing, accountancy)

Other

Vocational or work-related qualifications, other qualifications achieved in England or Wales, qualifications achieved outside England or Wales (equivalent not stated or unknown)

In 2023, almost nine million people lived in Greater London, making it the most populated ceremonial county in England. The West Midlands Metropolitan County, which contains the large city of Birmingham, was the second-largest county at 2.98 million inhabitants, followed by Greater Manchester and then West Yorkshire with populations of 2.95 million and 2.4 million, respectively. Kent, Essex, and Hampshire were the three next-largest counties in terms of population, each with around 1.89 million people. A patchwork of regions England is just one of the four countries that compose the United Kingdom of Great Britain and Northern Ireland, with England, Scotland and Wales making up Great Britain. England is therefore not to be confused with Great Britain or the United Kingdom as a whole. Within England, the next subdivisions are the nine regions of England, containing various smaller units such as unitary authorities, metropolitan counties and non-metropolitan districts. The counties in this statistic, however, are based on the ceremonial counties of England as defined by the Lieutenancies Act of 1997. Regions of Scotland, Wales, and Northern Ireland Like England, the other countries of the United Kingdom have their own regional subdivisions, although with some different terminology. Scotland’s subdivisions are council areas, while Wales has unitary authorities, and Northern Ireland has local government districts. As of 2022, the most-populated Scottish council area was Glasgow City, with over 622,000 inhabitants. In Wales, Cardiff had the largest population among its unitary authorities, and in Northern Ireland, Belfast was the local government area with the most people living there.

In 2023, the population of the United Kingdom reached 68.3 million, compared with 67.6 million in 2022. The UK population has more than doubled since 1871 when just under 31.5 million lived in the UK and has grown by around 8.2 million since the start of the twenty-first century. For most of the twentieth century, the UK population steadily increased, with two noticeable drops in population occurring during World War One (1914-1918) and in World War Two (1939-1945). Demographic trends in postwar Britain After World War Two, Britain and many other countries in the Western world experienced a 'baby boom,' with a postwar peak of 1.02 million live births in 1947. Although the number of births fell between 1948 and 1955, they increased again between the mid-1950s and mid-1960s, with more than one million people born in 1964. Since 1964, however, the UK birth rate has fallen from 18.8 births per 1,000 people to a low of just 10.2 in 2020. As a result, the UK population has gotten significantly older, with the country's median age increasing from 37.9 years in 2001 to 40.7 years in 2022. What are the most populated areas of the UK? The vast majority of people in the UK live in England, which had a population of 57.7 million people in 2023. By comparison, Scotland, Wales, and Northern Ireland had populations of 5.44 million, 3.13 million, and 1.9 million, respectively. Within England, South East England had the largest population, at over 9.38 million, followed by the UK's vast capital city of London, at 8.8 million. London is far larger than any other UK city in terms of urban agglomeration, with just four other cities; Manchester, Birmingham, Leeds, and Glasgow, boasting populations that exceed one million people.

Census 2021 data on religion by highest qualification level, by sex, by age, England and Wales combined. This dataset provides Census 2021 estimates that classify usual residents in England and Wales by ethnic group. The estimates are as at Census Day, 21 March 2021.

The religion people connect or identify with (their religious affiliation), whether or not they practise or have belief in it.
This question was voluntary and the variable includes people who answered the question, including “No religion”, alongside those who chose not to answer this question.

Total counts for some population groups may not match between published tables. This is to protect the confidentiality of individuals' data. Population counts have been rounded to the nearest 5 and any counts below 10 are suppressed, this is signified by a 'c' in the data tables.

This dataset shows population counts for usual residents aged 16 years and over. Some people aged 16 years old will not have completed key stage 4 yet on census day, and so did not have the opportunity to record any qualifications on the census.

These estimates are not comparable to Department of Education figures on highest level of attainment because they include qualifications obtained outside England and Wales.

Quality notes can be found here

Quality information about Education can be found here

Religion

The 8 ‘tickbox’ religious groups are as follows:

• Buddhist
• Christian
• Hindu
• Jewish
• Muslim
• No religion
• Sikh
• Other religion

No qualifications

No qualifications

Level 1

Level 1 and entry level qualifications: 1 to 4 GCSEs grade A* to C , Any GCSEs at other grades, O levels or CSEs (any grades), 1 AS level, NVQ level 1, Foundation GNVQ, Basic or Essential Skills

Level 2

5 or more GCSEs (A* to C or 9 to 4), O levels (passes), CSEs (grade 1), School Certification, 1 A level, 2 to 3 AS levels, VCEs, Intermediate or Higher Diploma, Welsh Baccalaureate Intermediate Diploma, NVQ level 2, Intermediate GNVQ, City and Guilds Craft, BTEC First or General Diploma, RSA Diploma

Apprenticeship

Apprenticeship

Level 3

2 or more A levels or VCEs, 4 or more AS levels, Higher School Certificate, Progression or Advanced Diploma, Welsh Baccalaureate Advance Diploma, NVQ level 3; Advanced GNVQ, City and Guilds Advanced Craft, ONC, OND, BTEC National, RSA Advanced Diploma

Level 4 +

Degree (BA, BSc), higher degree (MA, PhD, PGCE), NVQ level 4 to 5, HNC, HND, RSA Higher Diploma, BTEC Higher level, professional qualifications (for example, teaching, nursing, accountancy)

Other

Vocational or work-related qualifications, other qualifications achieved in England or Wales, qualifications achieved outside England or Wales (equivalent not stated or unknown)

Anthropogenic noise can adversely impact urban bird populations by interfering with vocal communication. Less research has addressed if anthropogenic noise masks the adventitious sounds that birds use to aid predator detection, which may lead to increased vigilance and reduced feeding efficiency. We test this hypothesis using a controlled playback experiment along an urban-rural gradient in Sheffield (UK). We also test the related predictions that anthropogenic noise has the greatest impacts on vigilance and feeding efficiency in rural populations, and on species that are more sensitive to urbanisation. We focus on six passerines, in order from most to least urbanised (based on how urbanisation influences population densities): blue tit Cyanistes caeruleus, robin Erithacus rubeculla, great tit Parus major, chaffinch Fringilla coelebs, coal tit Periparus ater and nuthatch Sitta europaea. We used play-back of anthropogenic urban noise and a control treatment at 46 feeding stations located along the urban-rural gradient. We assess impacts on willingness to visit feeders, feeding and vigilance rates. Exposure to anthropogenic noise reduced visit rates to supplementary feeding stations, reduced feeding rates and increased vigilance. Birds at more urban sites exhibit less marked treatment induced reductions in feeding rates, suggesting that urban populations may be partially habituated or adapted to noisy environments. There was no evidence, however, that more urbanised species were less sensitive to the impacts of noise on any response variable. Our results support the adventitious sound masking hypothesis. Urban noise may thus interfere with the ability of birds to detect predators, reducing their willingness to use food rich environments and increase vigilance rates resulting in reduced feeding rates. These adverse impacts may compromise the quality of otherwise suitable foraging habitats in noisy urban areas. They are likely to be widespread as they arise in a range of species including common urban birds.

Methods The methodology used to collect these data is described in Merrall, E. S. and Evans, K. L. 2020. Anthropogenic noise reduces avian feeding efficiency and increases vig-ilance along an urban-rural gradient regardless of species’ tolerances to urbanisation. – J. Avian Biol. 2020: e02341

Site selection and urbanisation metrics

Work was conducted in and around Sheffield (53°22′N, 1°20′W), which is England’s fifth largest city, with a population of c. 575,000. We defined urban areas as 1km x 1km grid cells with more than 25% impervious surface (following Gaston et al. 2005) and selected 16 urban sites within 3 km of the city centre, 19 urban sites that were more than 3 km from the city centre and 11 rural sites that were between 1 and 3 kms away from the city’s outer limits (using the above definition of urban areas). This approach enabled us to haphazardly select sites that were spread across the urban to rural gradient (see below for quantification), with the fewest sites in rural areas due to their greater homogeneity in background noise levels. All sites within the urban area were public parks and green-spaces with some woody vegetation cover, and all rural sites were woodland (rather than the alternatives of moorland or farmland) to maximise the similarity of the focal bird community along the urban to rural gradient.

Urbanisation intensity was quantified at each site using the ‘Urbanisation Index’ software developed by Seress et al., 2014 (available at: https://keplab.mik.uni-pannon.hu/en/urbanization-index). This software uses a semi-automated method where it takes a 1km2 area from google maps around the coordinates of each location, and then uses manually inputted training points to score each image for vegetation cover, forest, buildings and paved roads using 100x100m2 cells. It then uses principle component analysis (PCA) to calculate an urbanisation intensity score for each area.

Experimental design

Field work took place between the 12th February and 3nd April 2019. Our general approach was to assess avian feeding and vigilance rates at feeding stations under three experimental conditions: play back of urban noise and two control treatments (play back of natural noise and a silent control without playback). The urban recording was created by splicing together 5-minute sound recordings from each of four locations in central urban Sheffield using a Zoom H4n sound recorder and Cubase LE AI Elements 10, with a mixture of traffic, pedestrian and construction noise. The urban noise treatment was played at approximately 80 decibels (dbc), i.e. the typical volume of anthropogenic noise in busy urban areas during the day (Maryland SHA, 2018). The natural control used a mixture of songs of summer migrants (barn swallow Hirundo rustica; whinchat Saxicola rubetra; redstart Phoenicurus phoenicurus and common whitethroat Sylvia communis). Recordings were obtained from Xeno-canto (2005-2019) and were vetted to ensure that they did not include alarm calls to ensure that this treatment did not include vocalisations to which birds using the feeders were likely to respond. The natural sound control treatment was played at approximately 40 decibels (dbc) that matched natural sound levels of avian vocalisations. The urban treatment lasted 40 minutes while each control lasted 20 minutes, with a 10-minute habituation period between equipment set-up and beginning playback, and between the urban treatment and the two control treatments (which each lasted 20 minutes). We used two green, portable SONY SRS-XB10 Bluetooth controlled speakers (IPX rated 5) at each site located approximately a metre from the feeding station in a spatial configuration that created a surround sound effect.

Each site’s feeding station was set up four to seven days before conducting the experiment to enable birds to habituate to the presence of the food source. Each station consisted of two standardised hanging feeders, each with two feeding ports, filled with sunflower hearts. These are a nutritious food source that has a negligible handling time and is thus widely used by a wide range of species. To reduce the risk of disturbance from grey squirrels Sciurus carolinensis, which can discourage birds from using feeders (Bonnington et al. 2014), poles were greased and sunflower hearts were coated in chilli powder (which squirrels avoid, whilst birds exhibit no adverse response). Feeders were placed in relatively open locations to allow easy observation, but close to vegetation cover to encourage birds to approach and use the feeders. Feeding stations were located away from footpaths to minimise effects of human disturbance.

Treatments were applied in a randomised order at each site (with the three treatments being applied sequentially). Sites were visited in a haphazard manner with regard to the site’s urbanisation score. All data collection took place at least 1.5 hours after civil dawn and before civil dusk, to avoid spikes in bird activity early and late in the day. Data were not collected when it was snowing or raining (except occasional light drizzle), or at high wind speeds as such conditions interfered with activity levels and noise transmission.

Each treatment was filmed using a Panasonic (HC-X920) HD Camcorder and observed from an approximately 10m to 15m distance. Videos recorded birds feeding on the feeder and those feeding on spilt food beneath the feeder. If disturbance events occurred during the treatment, such as a human or dog passing close to the feeder, and interrupted birds’ feeding behaviour, data collection was paused until birds resumed normal activity. Videos were analysed, and for each visit we recorded the species, visit duration (seconds), number of pecks (as a measure of feeding rate) and the amount of time (seconds) spent performing vigilance behaviour, defined following Quinn et al. (2006), as when the bird raised its head and scanned. For each site, the temperature (oC) and wind speed (kn) were also recorded using data from the nearest weather observation site (Met Office, 2019). The seven weather stations used ranged from 220 m to 6.1 km away from the study site.

Statistical analysis

All statistical analyses were conducted using R Studio (RStudio Team, 2016). Three response metrics were calculated from the videos for each species per treatment per site: i) visit rate, i.e. the rate at which the species visited the feeder per hour of treatment, ii) the peck rate per hour of treatment, and iii) the vigilance rate (in minutes) per hour of time spent on the feeder. A total of 19 species were observed using the feeders or feeding on fallen seed on the ground below, of which six occurred at ten or more sites and were included in data analysis (great tit Parus major 46 sites; blue tit Cyanistes caeruleus 44 sites; coal tit Periparus ater 33 sites; Eurasian robin Erithacus rubeculla 35 sites; nuthatch Sitta europaea 14 sites; and chaffinch Fringilla coelebs 13 sites). These species vary substantially in their ability to maintain high population densities in urban environments. This is indicated by their urbanisation scores as calculated by Evans et al. (2011), i.e. the ratio of urban to rural population densities obtained from Breeding Bird Survey data from approximately 3,000 randomly selected 1 km x 1 km squares located across the UK. These scores, from most to least urbanised species, are: blue tit 1.46; robin 0.99; great tit 0.74; chaffinch 0.25; coal tit 0.23, nuthatch 0.17. Matched paired t-tests demonstrated that each of these six species’ visit, peck and vigilance rates did not differ between the two forms of control (P ranges from 0.110 to 0.877) and these data were thus merged to form a single control treatment.

We modelled visit, peck and vigilance rates using mixed effect models (lme4 package; Bates et al, 2015). These models pool data across