During the National Rifle Association’s May 2016 convention, where Donald Trump won the organisation’s coveted endorsement, board member David Keene hosted a private dinner that drew three Russian lifetime NRA members: a since-convicted Kremlin influence agent, her handler and mentor, and a previously unreported Russian who chairs a defense industry foundation, the Guardian has learned.

For the powerful 5 million-member NRA, the Russia ties of Keene and other NRA bigwigs are political and legal headaches that won’t go away – not with the FBI and congressional panels pursuing lengthy probes into the Kremlin’s alleged scheme to influence the pro-gun giant’s top brass, other conservative groups and US politicians.

In fact, as American politics has become consumed with the impact of Russia’s attempt to disrupt the 2016 presidential election in favor of Donald Trump, the NRA and some top board members have also become ensnared in Russia-related investigations, delivering an unexpected blow as the NRA’s links to Russia come under increasingly close examination.

The links are a complex web as evidenced by two of those at the 2016 convention dinner. Ex-Moscow banker Alexander Torshin, who was sanctioned last year, and his gun rights protege Maria Butina, who in late 2018 pleaded guilty to being a Russian influence agent and is now cooperating with US law enforcement, spent years cozying up to the NRA and some of its leaders. Their goal was to “establish unofficial lines of communication with Americans having power and influence over US politics”, according to Butina’s later plea agreement.

US prosecutors, who charged Butina with conspiracy to act as an agent of Russia, have indicated that GOP operative Paul Erickson, who worked closely with her, is a target too, and Torshin was her handler.

Moreover, Senate and House panels are also continuing probes that began in 2018 into the NRA’s Russia ties, and are looking harder at what NRA leaders knew about Butina and Torshin’s real goals, and a controversial trip they hosted in late 2015 in Moscow for several NRA honchos.

“The public record is clear that the NRA bent over backwards to help a Russian agent insinuate herself into conservative political circles,” Senator Ron Wyden, a member of the intelligence and finance committees, said in a statement. “The public has a right to know who at the NRA knew Torshin and Butina’s agenda, and why they were so eager to help these Russian agents.”

NRA chief Wayne LaPierre. Photograph: Chip Somodevilla/Getty Images

Questions about the NRA’s Moscow ties and the gun group’s role in the 2016 elections were also spurred when McClatchy reported last year that the FBI was looking into allegations that Torshin and other Russians may have improperly funneled Russian funds into NRA coffers as part of its record setting $30m in pro-Trump spending.

The NRA has stated it did not spend any foreign funds in the elections, which would be illegal, and said it only received about $2,500 from Russian sources for non-electoral purposes in the 2016 cycle.

Still, Russia intelligence veterans say Torshin’s work with Butina bears hallmarks of a Kremlin operation that targeted the NRA – and had some success.

“It’s most likely that the Kremlin reached out to Torshin or he went to them” about his NRA cultivation efforts, said Steven Hall, a retired CIA chief of Russia operations. “But either way, Torshin used Butina as part of the Kremlin’s broader influence efforts.”

Butina and Torshin’s long cultivation of NRA leaders involved attending several NRA conventions, and wining and dining a group of NRA bigwigs including, board member Keene, who visited Moscow in late 2015 for a week of meetings with top Russian officials organized by the Butina-led gun group, Right to Bear Arms.

Butina actually began working closely in 2015 with Erickson, an NRA member and fundraiser, who facilitated her entree into Republican circles nationwide and was romantically involved with Butina.

The charging documents against Butina cite her efforts to influence an unnamed “gun rights organization”, which is said to be the NRA. Although Erickson and Torshin are not named in court filings, the descriptions of the two individuals who aided Butina match the roles that Erickson and Torshin played in her Kremlin-blessed influence operations.

The NRA’s Russia ties have drawn scrutiny in part because some of Butina and Torshin’s contacts with the NRA and the Trump campaign have echoes of the Kremlin’s broader operations to help Trump become president.

Some close NRA allies say the Butina-Torshin efforts to woo the NRA was fueled heavily by Moscow’s interest in boosting Russian gun exports and easing sanctions, a key Putin goal.

“It’s smart to follow the money,” said Joe Tartaro, the president of the Second Amendment Foundation and an NRA ally, that sent some officials to an inaugural event for Butina’s group in 2013 that Erickson and Keene attended.

“I think people in Russia tried to cozy up to the NRA as much as they could,” added Tartaro. “They were trying to open US markets. They were interested in export business, not just for arms but ammunition as well.”

Further, emails from Erickson indicate that Torshin and Butina around the time of the 2016 NRA convention also tried to promote a meeting between Trump and Putin.

The NRA not only spent a record $30m to help Trump, but gave him an earlier than usual endorsement at their May 2016 convention and critical early ad support before his campaign’s ad blitzes.

Besides Torshin and Butina, the other Russian who dined with Keene at the NRA meeting was Dmitriy Osipkin, who chairs the Foundation for the Development of Promising Defense Technologies, according to two sources familiar with the dinner.

That evening Donald Trump Jr was dining with a larger NRA contingent and others at the same restaurant, enabling Torshin to chat briefly with Trump’s son.

These and other Russia linkages with the NRA and the Trump campaign seem to explain why special counsel Robert Mueller’s probe of Russian meddling in the 2016 elections has looked at the ties between the NRA and the Trump campaign, according to CNN.

Torshin’s contacts with the NRA began around 2011 when he attended his first NRA convention; two years later Keene and Erickson were in Moscow for a Right to Bear Arms meeting hosted by Butina and Torshin.

In December 2015, Torshin and Butina invited an elite NRA group to Moscow where they were feted at vodka and caviar dinners. On the trip were incoming NRA president Pete Brownell, NRA fundraiser Joe Gregory who runs a program for donors who kick in $1m a year, and Keene.

The NRA’s Moscow itinerary featured meetings with the foreign minister, Sergei Lavrov, and the deputy prime minister, Dmitry Rogozin, who oversaw defense and munitions industries.

Maria Butina, who in late 2018 pleaded guilty to being a Russian influence agent and is now cooperating with US law enforcement, spent years cozying up to the NRA. Photograph: AP

Key congressional committees and Democrats have been particularly interested in learning more details about the NRA Moscow trip.

Last December, Wyden requested documents from and interviews with Keene, Brownell and ex-NRA president Allan Cors, who opted not to go on the Moscow trip in late 2015 after the NRA’s top executive Wayne LaPierre raised concerns with him about participating, Cors said in a statement. None have agreed to be interviewed by Wyden’s office, but Brownell has provided documents, said a source familiar with the requests.

A Democratic aide with the House intelligence committee said in a statement that it expects to look more closely at the NRA’s Russia links and “whether and to what extent Russians sought to use the NRA as a back channel and the allegations that Russians may have funneled money through the organization to influence the election”.

The NRA and other not-for-profits can accept foreign donations for non-electoral purposes, but it’s illegal for foreign funds to be spent on elections.

NRA outside counsel William Brewer III said the NRA hasn’t found any evidence that Russian funds were spent improperly in the elections.

“The NRA believes that no foreign money made its way into the organization for use in the 2016 presidential election,” Brewer said in a statement. The NRA “has controls in place to vet contributions to the organization”.

But Wyden and others in Congress remain generally skeptical of the NRA’s statements about its Russia links, and is seeking more financial records to verify the scope of their activities.

“The NRA has offered many different stories about its relationship with Russia,” Wyden added, stressing that “no one should take their statements at face value”.