Joss Whedon, lauded king of fandom, purveyor of wisdom and de facto overseer of the Marvel Cinematic Universe, celebrates his 50th birthday on Monday (June 23).

Whedon's always had a knack for taking pre-established characters – whether it's The Avengers or a gang of talking toys or the cast of Speed – and making them just that bit more quippy and emotionally rich and awesome. But it's his original small-screen characters that remain his most beloved.

Below, Digital Spy counts down our top fifteen Whedonverse characters. You can vote for your favourite in the poll below, then take to the comments to rank about all the rankings we got wrong and all the favourites we've inevitably left out.

15. Dr Horrible



Neil Patrick Harris's "aspiring supervillain" is never at any point successfully villainous. He's a more immediately loveable extension of Buffy's Nerd Troika – the perpetually overlooked semi-recluse who can barely string a sentence together around his crush, and thinks that evil is the way to finally get noticed.

"The world is a mess, and I just need to rule it," Dr Horrible reasons, but he really doesn't have the stomach for the kind of wrongdoing that will get him his coveted spot in the League of Evil. His misanthropy is only bolstered as Penny, the long-time object of his affections, becomes infatuated with Nathan Fillion's pompous, smarmy superhero Captain Hammer.

Even as things go tragicomically wrong for Dr Horrible in classic Whedon fashion, Harris remains a totally endearing central presence. He's also kind of deep, like when he tries to warn Penny that layered people aren't always good news: "Sometimes there's a third, deeper level that's exactly the same as the surface one. Like pie."

14. Cordelia Chase



If there's one thing Whedon excels at, it's taking a character from zero to hero. Cordelia started life in Buffy's very first episode as the archetypal high school mean girl, a bitchy thorn in the side of the Scooby Gang. The best thing you could say about her was that she was brutally honest, though more often just brutal.

But her blunt self-interest made her brave and resourceful, and she rapidly became a reluctant part of the team - all the more so after losing her status when she dated Xander and, even worse, fell for him. By the time she ends up penniless in Los Angeles, she's a natural fit for Angel's fledgling detective agency.

While Angel's difficult fourth season saw the character screwed over by dodgy writing (and behind-the-scenes drama, if the grapevine is to be believed), she's still one of the Buffyverse's most dynamic characters, and her final send-off in season five's 'You're Welcome' is an understated tearjerker.

13. Andrew Wells



Season six is nobody's favourite run of Buffy, and the shambolic Nerd Trio were nobody's favourite villains, though their ineptitude did make for a welcome contrast early on to the season's generally dark tone. While ringleader Warren was a one-note bully and misogynist, his de facto right hand man Andrew was essentially sweet but fatally weak-minded.

It's fair to say Andrew represented a fair chunk of the audience, with his incessant pop culture references and preference for living in a fantasy world over reality, and while he never quite made the zero to hero transition on screen, he is one of the few human characters to get a fully-fledged redemption arc. Season seven's Andrew-centric 'Storyteller' is a witty and touching highlight of an uneven final season, while Angel's much stronger last episodes still benefit from the brief reappearance of Andrew and his crush on Spike.

Providing comic asides and emotional texture without drawing attention away from the main cast, Andrew did everything a good supporting player should.

12. The Mayor



Buffy's second-best villain (more on which below) is also one of its simplest. There's no intricate backstory or hidden complexities with Mayor Richard Wilkins – he's just a jovial family man who dislikes swearing, values his work, and ultimately plans to transform himself into a giant snake demon. As gleefully played by Harry Groener, he's a feel-good sitcom dad with occasional flashes of pure menace.

And while there's wisely no attempt to humanise or redeem the Mayor, what does stand out is his twisted father-daughter relationship with Faith, whose volatility and desperate need for guidance make her an ideal blunt instrument for the Mayor to exploit.

11. Kaylee Frye



"I don't believe there's a power in the 'verse that can stop Kaylee from being cheerful – sometimes you just want to duct tape her mouth and dump her in the hold for a month," Mal says fondly of Serenity's mechanic. It's a testament to Jewel Staite that the audience never shares his feigned irritation – relentlessly sunny characters are tricky to get right, particularly in a universe that generally tends towards cynicism.

But that's exactly why Kaylee is such a standout. Firefly is a more morally grey show than either Buffy or Angel, centred as it is on a crew of smugglers, thieves and fugitives, and she's the sweet, open-hearted calm at the eye of the storm.

10. Angelus



Buffy's central "high school is literally hell" metaphor was never explored more effectively than in the moment where Buffy, having lost her virginity to brooding, sensitive vampire boyfriend Angel, wakes up to find him gone. Later, she's relieved to find him back at his apartment, only to find that he's transformed into somebody she barely recognises – somebody cruel and disinterested.

"You've got a lot to learn about men, kiddo," he smirks, pityingly. "Although I guess you proved that last night." It's a truly brutal moment that rings true in the way that all Whedon's best writing does; the supernatural element of the storyline is close to irrelevant, because the emotional reality is so painfully recognisable to anyone who's ever been a teenager. Angelus is the best villain Buffy ever had, because the threat he posed and the ammunition he carried was so personal.

While Boreanaz took a good two seasons to really grow into the character of Angel, he seemed immediately at home with the pure, caustic savagery of Angelus, and his genuinely frightening blackened dynamic with Buffy is a huge part of what makes season two arguably the best of Buffy's entire run.

9. Jayne Cobb



Jayne is a pretty terrible human being. From the outset, Serenity's resident gun-for-hire is established as aggressive, venal and crass – one of his very first lines of dialogue is a crude dig at Kaylee for getting "lubed up" over Simon, and hurting Kaylee is generally shorthand for pure evil in the Firefly universe.

He also makes it clear that he'll willingly turn on Mal and the crew if the price is right, a promise that he follows through on when he sells Simon and River out to The Alliance in 'Ariel'. Mal very nearly throws him out of the airlock for this, and he deserves it.

And yet between Adam Baldwin's brash, buoyant performance and the consistently quotable lines he's given, Jayne quietly becomes loveable despite all the odds. Had Firefly been given the five-season run it so richly deserved, he'd undoubtedly have got a lot more complex. As it is he's just unexpectedly fun, from his willingness to wear headgear knitted by his mother, to naming his gun Vera, to his iconic one-liners.

8. Anya Jenkins

20th Century Fox



Another intended guest star who proved herself indispensable was Emma Caulfield as demon-turned-human Anya. Formerly a vengeance demon who specialised in punishing unfaithful men, she's forced to endure the indignity of living life as a human, and the even worse indignity of her growing feelings for the initially reluctant Xander.

Anya's staggering bluntness was a consistent goldmine for comedy, with her character filing a similar role in the Scooby Gang to the one vacated by Cordelia. But in episodes like 'The Body', in which she struggles to come to terms with the reality of death, it's also immensely touching, and the growing humanity beneath Anya's prickly surface made her one of Buffy's most unexpectedly tragic, loveable characters.

7. Willow Rosenberg



No character in Buffy goes unchanged, but the transformation of Alyson Hannigan's Willow from meek, overlooked wallflower to scarily self-possessed makes her easily the most dynamic of the core players.

The early potential for Dark Willow is explored in two of the show's best episodes of all time – 'The Wish', which portrays an apocalyptic future in which Willow is a vampire, and 'Doppelgangland', in which vampire Willow comes to regular Sunnydale – and the contrast worked so well that Willow herself gradually moved in a darker direction as she grew unhealthily dependent on her magic, unable to maintain any confidence without it. While many viewers were left feeling nostalgic for the loveable, quirky Willow of the early seasons, her progression is a testament to how disinterested Whedon and his co-writers always were in playing it safe.

Willow was also a touchstone for Buffy's broader cultural significance, being one of the first major TV characters to have an openly gay relationship. Willow and Tara were treated in exactly the same way as any other romance on the show, and given how rare this still is in mainstream television, it feels all the more powerful in retrospect.

6. Rupert Giles



While Buffy had more than her fair share of complex, doomed romances, her most significant relationship of all might actually have been with mentor and surrogate father Giles.

Their early dynamic was one of affectionate push and pull, with Buffy still trying hard to be a peppy normal teenager and Giles at his most bookish and stern. But as Giles' dark past was revealed and Buffy gradually came to respect him (and her calling) more fully, they became the core of the show, occasionally spotlighted in standout episodes like 'Passion' and 'Helpless'.

Later on Anthony Stewart Head's knack for comedy (and spectacular singing voice) were exploited more and more in the post-season 3 years, when Giles was no longer a Watcher or a librarian, but did occasionally wear outfits like this or sing songs like this. Giles would be even higher on this list if it weren't for the strange semi-character assassination that occurred in season seven, and poisoned the Giles/Buffy relationship in a way that was never really resolved.

5. Faith Lehane

20th Century Fox



Given the apparent randomness of the Slayer "chosen one" process, it stands to reason that not every girl's cut out for the job. With the introduction of Faith in season three, Whedon deepened the mythology surrounding the Slayer, asking the question: what would the burden of a forced destiny and wildly enhanced physical abilities do to somebody who was already unstable?

Unlike Buffy, Faith's upbringing was chaotic and violent, and her approach to her mythic responsibility is similar. Eliza Dushku's performance is so seductive and affable that you're totally with Buffy when she starts shirking school to go and party with Faith, revelling in the chance to embrace her inner bad girl.

And then it all goes very bad, very fast, and Faith becomes one of the most complex, unpredictable and darkly entertaining characters in the Buffyverse, wreaking gleeful body-swapping havoc before finally getting her own messy, nuanced redemption arc on Angel. While the fascinating Buffy/Faith relationship never got the resolution it needed in season seven, the decision to bring her back for the end was a wise one. A moment of silence here for the long-mooted Faith spin-off series.

4. Malcolm Reynolds



While TV anti-heroes are now a dime a dozen, they were a much rarer breed when Firefly debuted in 2002. A roguish space cowboy with a fluid moral code and a deep-buried heart of gold, Mal still looks like an outright hero when stacked against the Walter Whites of the world, but at the time he was pretty revolutionary as a leading man.

So much so that Fox asked Whedon to lighten the character up from the show's famously brilliant, famously unaired pilot 'Serenity', making him more quippy and more charmingly crotchety as opposed to emotionally closed off. Whedon was largely able to blend the two without much of a contradiction – Nathan Fillion is a naturally winning and witty performer, even when playing a man whose faith has been shattered by war and corrupt government.

At risk of sounding like a broken record, it's almost physically painful to imagine just how much more fascinating Mal would have become had Firefly survived. As it is, his understated wisecracking, emotionally complexity and uniquely fierce brand of loyalty still make him top five material.

3. Spike

20th Century Fox



Spike was everything throughout the course of Buffy – punk rock villain, comedy sidekick, lovelorn anti-hero, vicious stalker, reformed champion – and this kind of rogue element could very easily have worn thin. Instead, he became one of the show's major breakout characters, largely because James Marsters's performance provided such a strong core that the writers could get away with spinning just about anything around him.

The intriguing thing about Spike is that he broke the rules that had been established with the Angel/Angelus split. Even without a soul, he was capable of deep love and even of friendship, which made the show's supernatural universe far more morally complex and knotty.

While the long-built Buffy/Spike relationship was fumbled somewhat in the gratuitously bleak sixth season (Whedon's absence as head writer being noticeable), Spike's slow, uneasy journey to redemption was utterly compelling. And after the heavy emphasis on his angst in latter day Buffy, Spike's reincarnation on Angel as a reluctant sidekick and snarky thorn in Angel's side was a reminder of just how funny the character could be.

2. Wesley Wyndham-Pryce



Other characters might have lost their souls and regained their souls and returned from the grave, but it was Watcher-turned-rogue demon hunter Wesley who ended up changing the most dramatically out of everybody in the Buffyverse, with Willow being his only close rival.

Introduced in Buffy's third season as a wimpy counterpoint to the increasingly badass Giles, Wesley was good as a punchline and very little else. When fledgling spinoff Angel found itself in need of a new series regular after the unfortunate departure of Glenn Quinn, Wesley wasn't the likeliest candidate.

While he's still used largely for comedy in Angel's early going (his hammy imitation of Angel alongside Cordelia's Buffy remains a series highlight), there was a subtle shift to laughing with him, rather than at him. Before long he becomes a loved and trusted part of the Angel Investigations team, and a central character. Which, this being a Joss Whedon show, translates to suffering. Lots of suffering.

After a false prophecy led Wesley down a dark path in season three and four, it became clear just what a compelling dramatic actor Whedon had in Alexis Denisof. Even once Wesley returned to the fold at Wolfram & Hart, things only got bleaker and Denisof's performance only grew more mesmerising. In perhaps the most brutal example of Whedon's self-confessed cruel streak, Wesley's long-time love Fred died a slow death in his arms just days after the pair finally got together.

The hollowed-out Wesley of Angel's final episodes is almost unrecognisable from where he started, and yet both incarnations of the character feel lived-in and authentic – it's because the transformation is so subtle that the contrast is startling.

1. Buffy Summers



Whedon initially conceived his Slayer as an answer to horror cinema's cliché of the doomed, tiny blonde girl alone in a dark alleyway. But an inverted stereotype isn't enough to build a show around, no matter how empowering, and what Sarah Michelle Gellar's Buffy represents very quickly became secondary to what she is.

She's first and foremost a teenage girl, albeit one with bravery and wisdom way beyond her years, and Gellar's heartfelt, vibrant performance made even her most otherworldly experiences feel universal. While she's no rebel compared to Faith, she does question her calling and crucially rejects the Slayer doctrine of working alone.

But the great thing about Whedon's writing is that he consistently gave Buffy much more immediate, human things to worry about than her destiny. Sure, she had to contend with ancient evil and with the apocalypse and with death (twice). But she also had her heart broken, and worried about her future, and struggled to adjust to college, and got used for sex by a terrible guy, and lost her mother suddenly to a brain aneurysm.

And while she did save the world a lot, she didn't do it without personal cost – Buffy ages and hardens a lot over the course of the show, and by season seven she's a shell of her former self. Nevertheless, she's still one of the most fully developed, enduringly human characters in TV history, a modern-day icon as beloved as any comic book superhero.

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