Despite reporting strong revenue figures for the last quarter, Twitter says it lost 4 million users and has Apple to blame.

During the company's quarterly earnings call with shareholders on Thursday, Twitter chief financial officer Anthony Noto announced that Twitter has 288 million monthly active users (MAUs), which is 7 million users short of the analyst forecast of 295 million monthly active users.

The culprit, said Noto, is Apple's iOS 8, which Cupertino began rolling out of its doors in September. Three million of those users lost were blamed on what Twitter calls auto-polling, while the remaining 1 million were "Twitter-owned."

"We don't expect to get the three million auto-polling MAUs in Safari back, and that's a non-Twitter owned and operated auto-polling MAU," Noto said during the call. "The one million, that number was actually higher at a different point in the quarter and we were able to bring it back to just one."

Prior to iOS 8, users of Apple's Safari browser had a Shared Links tab in its bookmarks section that include links to tweets of people that users follow on Twitter. In iOS 7, Safari linked to the user's Twitter account to ask for tweets to add in Shared Links. However, Apple discontinued this feature in iOS 8, causing Twitter to lose 3 million users who were presumably accessing part of their Twitter timelines from Safari.

The keyword here is "presumably," since auto-polling figures are an unreliable metric of how many people actively use Twitter every month. In iOS 8, users will still be able to receive links from Twitter, but only if they explicitly ask Safari to do so. This could give a more accurate picture of Twitter's active monthly user base than one that relies on automatic polling.

As for the other 1 million users lost, Twitter says it was due to an "unforeseen bug in the release of iOS 8 as it relates to the specific Twitter integration into iOS." Twitter CEO Dick Costolo tells BusinessInsider in a follow-up call that the problem was an encryption issue that prevented some users from logging in to their Twitter account on their iPhones and iPads.

"That was a much more complex issue, it did not have a one-size-fits-all fix, so the team here worked as quickly as possible to address it but it caused a large number of users to not be able to use the product, even those who were trying repeatedly to figure out ways to get in," Costolo says.  

Costolo took pains to emphasize that "we obviously have a great relationship with Apple."

Although Twitter posted a revenue of $479.1 million, a 97 percent increase from last year's figure, and reported a smaller net loss of $125.4 million, the microblogging network's monthly active user base is considered a key metric used by investors to evaluate Twitter's ability to monetize its social network by reaching wider audiences and earning more from advertising.

Twitter's shares were up 3.74 percent to $45 in after-hours trading following the call.

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