Sonic: Lost World pays homage to the Sonic series' past with a bevy of easter eggs — from appearances of old enemy types to familiar environmental design elements — while still keeping it fresh and relevant through updated game mechanics.

During hands-on time with the game, Sonic brand manager Aaron Webber and producer Sam Mullen told Polygon that Lost World features bits and pieces from previous Sonic titles, including bouncing mushroom enemies from Sonic and Knuckles, artwork from Sonic 2 featured in a casino area's spinning roulette wheels and appearances from a dozen other characters including Knuckles and Amy. The game also makes liberal use of Wisps from Sonic Colors, tiny creatures that give Sonic temporary powers like flying or being able to drill through the ground.

"Every enemy you see is a reference to a previous Sonic game, which could date all the way back to Sonic 1," Webber said. "So everything from the old Sonic games up until now make an appearance."

What separates Lost World's referential treatment from the nostalgia-driven Sonic Generations is that while the easter eggs and past references play a huge part in gameplay, they're not the main focus. The Sonic team wanted to create a world that was "100 percent Sonic" without leaning heavily on old mechanics and story beats. For example, for the first time in the series' history, Sonic is teaming up with his arch-nemesis Dr. Eggman to defeat an even greater threat to the both of them: new enemies called the Deadly Six. This rivalry "got kind of stale," Webber said, and Sega felt the leap to a new console was the perfect time to twist the story. This formula shakeup and sprinkling of familiar elements is a way to garner new interest in the series, and could encourage players to seek out previous games' based on Lost World's references — as well as provide a treat for series' veterans.

"In past Sonic games, Sonic would move really fast and he was kind of uncontrollable, and then you'd run into objects and just stop."

Among this mix of new and old is a jungle level lifted from Sonic and Knuckles, but with a twist to accommodate the game's new parkour system. Players will slide along rails and zip across narrow strips of grass avoiding swathes of muddy water infested with alligators that can deplete all of Sonic's gold rings in one chomp.

The game's parkour system was driven by the team's desire to make Sonic more fluid to control, and give series newcomers the chance to breeze along as quickly as longtime Sonic gamers.

"The whole parkour system, conceptually, comes from the simple fact that in past Sonic games, Sonic would move really fast and he was kind of uncontrollable, and then you'd run into objects and just stop," Mullen said. "So we went back to basic controls and said, 'Okay, how to we get past this? People don't like running around fast, fast, fast and then stop.' It started out with really simple things, like when Sonic hit a corner he would just glance past it. So we thought, what if he runs up walls or grabs ledges? And it sort of evolved from there into the state it is now.

"It all boils down to the fact that we want the player to maintain a flow, even if they're not having a sort of perfect playthrough," he added. "This way, the flow of a high-level Sonic player can be enjoyed even by a novice player."

Sega has also taken care to keep Sonic veterans from being bogged down with tutorials; in both the 3DS and Wii U versions of the game, the tutorial is completely optional, but on the Wii U the tutorial selection bubble has been removed to the top of the GamePad screen. In a pinch, players can select it at any time for tips. On the 3DS, players can run Sonic into floating question marks peppering each level for notes on how to scale certain obstacles.

Each stage also features a certain number of rare red rings. Collecting all of these, Webber said, will unlock "something special" that "Sonic fans are sure to love." What this surprise might be was not revealed, but Webber did confirm that the surprise is not the announcement of the third game in Sega's Sonic deal with Nintendo.

The most notable change to Sonic for Lost World is that players have more control over the blue hedgehog than in previous games. In addition to the new parkour system that allows Sonic to scale obstacles and run up walls, different enemy types will require different moves to take them down; the days of button mashing to jump on everyone's heads are over.

Sonic will need to jump, kick, homing attack and perform a number of tricky moves to knock giant spiders off their webs and expose huge caterpillars' vulnerable points in order to do them in for good. There's a decent handful of different enemy types in each area, and while the number of enemies and moves can get frustrating to remember, Webber said this was done to add more strategy to the game and give players reasons to use attacks other than Sonic's classic jump.

"It's a matter of giving players reasons to use that kick attack versus the homing attack, and having enemies where you'll have to use a certain combination, such as the homing attack and then a kick," he said. "It's really nice to have that kind of control in a Sonic game."

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Some areas have mechanics unique to their levels. In a casino level where Sonic must navigate a brightly-lit corridor strewn with light rails that will electrify him on impact, players can collect silver chips strewn throughout the area. These chips can be cashed in at the end of each level for additional points and even allow players to progress to the next level more quickly. In a snow-themed level, Sonic rolls up into a giant snowball and must be navigated across thin floating platforms with a rolling mechanic similar to the Super Monkey Ball games.

"It's like getting to look at our piece from a different perspective and a different angle."

In the 3DS version, Sonic is running forward as in the Wii U version, a break from the traditional 2D side-scrolling nature of most handheld Sonic titles. But each level in the Wii U version has been completely redesigned for the 3DS version, offering players two alternate experiences of the same game. Players have the same fluid control over Sonic and his bank of moves, but in areas that are "entirely different" from what they may have seen in the Wii U version.

"We're working with [Osaka studio] Dimps on the 3DS version, and Dimps has done a really great job on the previous handheld games," Webber explained. "But in particular with this one, it's like getting to look at our piece from a different perspective and a different angle. So it's the same style and look of the game, but it's as if someone else is giving their own take on it.

"We also wanted to give fans something brand new, so if you buy the Wii U version and then the 3DS [version], we didn't want to give you the exact same game," he added. "You beat those levels already. So for Sonic fans that are going to buy both, or even for people that will just get one or the other, you want something that not only works but feels at home on the platform. So 3DS will have some changes to it that make it feel more natural, better on the 3DS. And likewise there's some stuff on the Wii U that just makes more sense and fits better there."

Bringing the 22-year-old franchise into the next generation of gaming — with an audience that has a whole new set of standards and expectations — has been a challenge for Sega, but one they were ready to embrace.

"Sonic has never stood still."

"Sonic has never stood still — it isn't about churning out yesteryear's game with new levels," Mullen said. "That's not what Sonic is about. We did that with Sonic: Generations, but with Generations, that was the whole concept of the game. But Sonic as a property is about trying new things and taking him to places he's never gone before, and having him do new things.

"Lost World is all about going back and rethinking the way he controls, trying new things and just like we did with Colors [and the color powers]... Well, if these things work, you'll see them in future games," he explained.

As for the Nintendo-exclusive deal, Webber and Mullen said that the partnership "just made sense."

"If we trace Sonic's third-party roots all the way back to just after the Dreamcast, we released Sonic Adventure 2: Battle on the GameCube and it did phenomenally well," Webber said. "We think that's because there was a lot of overlap between the Sonic/Sega and Nintendo crowd at that time, and we just found historically that Sonic performs really, really well on Nintendo platforms. Sonic Colors did great, Sonic and the Black Knight did well and Sonic and Secret Rings did well — all Nintendo. When we look at the numbers, we thought — this is clearly where our audience is."

But as far as the blue hedgehog's possible appearance in the upcoming Super Smash Bros. Wii U?

"We have no idea!" Webber said.