Harry and Meghan passed another royal milestone as husband and wife when they joined her in acknowledging the crowds gathered in the Mall following the traditional ceremony to mark her official birthday.
The Queen was flanked by her sons, the Prince of Wales and Duke of York, at the front of the balcony, while the newlyweds stood behind her, surrounded by other members of the royal family.
The younger royals lined up in front of the adults to enjoy the fly-past of modern and historic aircraft, with Prince George and Princess Charlotte standing with their parents, the Duke and Duchess of Cambridge.
Charlotte clapped excitedly when the thousands of people gathered at the gates of Buckingham Palace applauded the aerial display.
But Peter and Autumn Phillips’ seven-year-old daughter Savannah, who was with her sister Isla, could be seen playing up as everyone stood still while the national anthem was played.
The youngster could be seen fooling about with George and jokingly put her hand over his mouth.
The Duke of Edinburgh, who has effectively retired from official public duties, was the only major figure missing from the gathering.
Other members of the royal family on the balcony included the Duchess of Cornwall, Princesses Beatrice and Eugenie, the Princess Royal, the Countess of Wessex and her children, Lady Louise and James, Viscount Severn, but the Earl of Wessex was not present as he is thought to be returning from an overseas trip.
Earlier, the pomp and pageantry of Trooping the Colour was staged on Horse Guards Parade - a first for Meghan.
The duchess and her husband enjoyed a carriage procession to the event with other members of the royal family before they all watched the spectacle from Wellington’s old office.
Meghan looked chic in a dress by Carolina Herrera and a hat by Philip Treacy, while Kate was stylish in an Alexander McQueen dress and a hat by Juliette Botterill.
Camilla wore a pale blue silk dress and coat by Bruce Oldfield, and a hat by Philip Treacy.
Four generations of the royal family were gathered on Buckingham Palace’s balcony to watch the traditional flypast which featured modern Typhoon jets and also aircraft from the RAF Battle of Britain Memorial Flight - an iconic Spitfire, Hurricane and Lancaster bomber.
Andrew took up a more prominent place on the balcony after taking part in his first Trooping the Colour ceremony on horseback in his new role as Colonel of the Grenadier Guards.
He joined the other royal colonels riding behind the Queen’s Ascot Landau carriage, Charles who is Colonel of the Welsh Guards, Anne, Colonel of the Blues and Royals, and William, Colonel of the Irish Guards.
The Trooping the Colour ceremony is an event the Queen knows well and she would have inspected the Guardsmen, in their scarlet tunics and bearskins with a keen eye.
Following her recent eye operation to remove a cataract, the Queen showed no ill effects and was not wearing sunglasses as she has at some outdoor events.
The ceremony is not only the official birthday celebration of the monarch, but a major social event for the families of the servicemen and women on duty on the parade ground.
They watched as this year’s colour, the flag of the 1st Battalion Coldstream Guards, was trooped on Horse Guards, Henry VIII’s former jousting yard.
In a lighter moment, a young fox was spotted on the parade ground before the spectacle began, running out from under the stands as it tried to find a way into nearby St James’s Park.
But later in the day, as the Queen’s procession made its way back to Buckingham Palace, Field Marshal Lord Guthrie, a former head of the armed forces, was thrown off his horse near the Victoria Memorial.
A Ministry of Defence spokesman said he was “receiving medical attention”.