So the traditional World Cup warm-up double header against England is over for another four years.

Wales have one more warm-up match, against world champions South Africa in Cardiff on 19 August, before Warren Gatland names his 33-man squad two days later.

Following the opening 20-9 win over England in Cardiff, what will Gatland have learned from the 19-17 defeat at Twickenham, that provided one of the most extraordinary final quarters of a match in recent times including a red card for Owen Farrell.

Gatland angry at loss

The Wales boss admitted he was furious at how his side managed the final quarter when they let a 17-9 lead slip from their grasp.

Wales built an eight-point advantage following an excellent Tomos Williams' try with England at one point down to 12 men with Farrell, Ellis Genge and Freddie Steward off the pitch.

Wales showed concerning naivety as they conceded a penalty straight from kick-off in their 22 and allowed Maro Itoje to score from a driving line-out, despite the hosts having three fewer men on the field.

Gatland's side had been clinical in Cardiff when they absorbed England's first-half pressure before pulling away to win.

The capitulation in Twickenham could not have been more contrasting and Gatland was visibly angry. He admitted it had told him a lot about some of his players with perhaps some World Cup places on the line.

He will hope that will prove a valuable lesson and one that will not be repeated in France when the important action begins in September.

Line-out woes

Wales' line-out was good in Cardiff. Functioning at least. At Twickenham it was abysmal as the visitors lost six line-outs with hookers Dewi Lake and Sam Parry regularly failing to find their men.

England targeted this area from the outset with a voice shouting 'Pressure on Lake, Pressure on Lake' from the opening set-piece as the new Wales captain was attempting to throw.

That pressure never eased and even the change of hooker in the first half, when Lake was forced off injured, did not alter fortunes.

The line-out has proved an Achilles heel for Wales at different times in the modern game. Gatland will want to ensure there is marked improvement in this facet before Wales face Fiji in Bordeaux.

Discipline will also be a concern with yellow cards for Tommy Reffell and Adam Beard, although England had three players sin-binned and Farrell red carded for a high tackle on Taine Basham.

Injury concerns

World Cup warm-up matches always throw up injury issues and this campaign is no different with hooker and lock proving the main concerns.

Lake was given the Wales captaincy at Twickenham but left the field after injuring his knee at a ruck and Wales are waiting on news of the severity of his problem.

Gatland also announced after the match Ryan Elias is expected to be on the sidelines for four weeks following a hamstring injury suffered against England in Cardiff and battling to be available for the World Cup opener on 10 September.

Wales will need three hookers for the final squad with Parry and Elliot Dee the other specialists in the provisional squad.

Six Nations captain Ken Owens withdrew with a back injury that will rule him out of at least the World Cup group stages.

Exeter lock Dafydd Jenkins is also expected to be out of action for a month, although he does not require surgery on his knee. That is a blow having established himself as a genuine contender to partner Will Rowlands in the starting line-up.

Swansea-born Taine Plumtree, making his first start at number eight after his debut in Cardiff the previous week, is the other main injury concern as he left Twickenham in a sling after damaging his shoulder.

Gatland is looking for other number eight options with Taulupe Faletau battling a calf injury that has ruled him out of the three World Cup warm-up games.

Backs Gareth Anscombe, Johnny Williams and Alex Cuthbert are other injury concerns.

No ordinary Joe

Scarlets centre Joe Roberts was the one new cap to start at Twickenham with replacements Keiran Williams and Kemsley Mathias also making debuts.

Roberts was over eager when he was caught out in defence early on after rushing out of the line but following that he enjoyed an assured game and put himself into the selection mix.

"Joe did well and grew into the game as it went on," said Gatland.

That was most evident with his classy outside break and well-timed pass for Tomos Williams to sprint clear and score.

Roberts's partnership with Nick Tompkins was Wales' 21st different starting centre duo in 41 matches as they continue strive to find the suitable midfield combination.

The pair are competing with Keiran Williams, Johnny Williams, Max Llewellyn, George North and Mason Grady for probably four centre spots in Wales' final World Cup reckoning.

Class is permanent

Wales full-back Liam Williams had not played a game since March but you would never have known that judging by his impressive Twickenham showing.

His assured defensive work repelled England throughout with expert positional play. The 32-year-old was also elusive in attack when the few opportunities arose and was always looking for work.

Keeping Williams fit has been the issue in recent years and he will be vital for any World Cup challenge.

Whether Gatland wants to take another specialist to France in Leigh Halfpenny remains to be seen with either four or five back-three players being included.

The alternative will be to rely on part-time full-backs like Louis Rees-Zammit, Josh Adams and Gareth Anscombe to help fill the number 15 jersey.

Another old head, Dan Biggar, demonstrated his experience with an impressive cameo as he came on for the ineffective Owen Williams.

His clever cross-kick set up the penalty try after Wales wing Josh Adams was taken out in the air by England full-back Freddie Steward and he created space for Joe Roberts in the lead up to the second try.

Springboks await

Gatland has given opportunities to 42 players in two very different teams for the England encounters.

The Wales coach indicated there are still some players he wants to give chances to against South Africa with the world champions in Cardiff next weekend.

Wing Cuthbert and centre Williams will be hoping to return from injury and have their own August audition.

With the World Cup opener just four weeks away, Gatland will hope to have a starting side that bears a degree of resemblance to the team that takes the field in Bordeaux next month.