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The 30 Best Games of 2021
By Paste Games Writers | November 30, 2021 | 9:00am
Games Lists best games
Despite being the first full year of the latest generation of game consoles, we're not going to remember 2021 for any technological breakthroughs. We're not going to remember it as a year of game design innovations or for the debut of some groundbreaking new series that will help define the current generation. If we remember 2021 for anything in the gaming industry, it'll be for matters outside the games themselves. We'll remember 2021 for the culture of sexual harassment and inequality that was exposed at companies as large as Activision Blizzard and Sony PlayStation, and as small as Fullbright. We'll remember 2021 for the supply chain issues that have made those new consoles difficult to find. We'll remember 2021 as the year where an end for the pandemic finally seemed close at hand, letting not just the games industry but life itself return to something like it used ot be, only for us to fumble the ball near the finish line. 2021 wasn't 2020, but it wasn't much better.
When we do actually consider the games of 2021, something notable sticks out. The best games of the year were new installments of older series rooted in older genres, only with a psychological and emotional depth we wouldn't normally expect from games of the past. Psychonauts 2 built on the emotional framework of the 2005 original and explored the memories, feelings, and regrets of its elderly characters. Ratchet & Clank: Rift Apart used its cartoon mascots to examine self-doubt, loneliness, and anxiety with charm, wit, and heart. Metroid Dread tapped into a vein of terror that gradually disappeared from the Metroid series over the decades, bringing it back with more power than before. These aren't the only memorable or moving games of the year, but they are the best, and it's because of the way they make us feel—not just the physical and intellectual satisfaction of playing them, but the way they touch our hearts and minds.
Yeah, I know that sounds schmaltzy. No regrets. It's alright to feel, y'know?
There are 27 other games on this list. They'll all make you feel many different things in many different ways. They're all worth playing, if you have the time and the necessary equipment. They all get the Paste Games stamp of approval, our nod of acceptance, a tip of whatever Georgia sports team hat we happen to be wearing that day. Good job, games.
Here are the 30 best games of 2021. Take a gander, if you'd like.
30. It Takes Two
Platforms: PlayStation 4, PlayStation 5, Xbox One, Xbox Series X|S, PC
Two-player co-op game It Takes Two's mundane settings are an opportunity to get wacky with mechanics and gameplay features, which the game will just fling at you. Every level explores a gimmick or series of gimmicks before casting it aside for the next, so it manages to stay remarkably fresh almost the entire way through. One second you're playing a shooter and the next you may be playing a hack and slash. I don't want to spoil my absolute favorite, but the amount of ludicrous things that come together to make it happen is nothing short of magic. Little of it makes any sense with or without context, but also It Takes Two comes across as a videogame for the sake of being a videogame, and while I respect that, it does mean the game shoots its own story in the foot often. The game's simultaneously asking you to care about this impending divorce and the effect it'll have on their daughter and the ludicrous task to gun down wasps or murdering plushies often! It forces the player to either try and reconcile these nonsensical aspects, or focus on a thing at a time. By the time I reached anything I've mentioned, I'd long since shut off my brain and decided to bask in the vibes rather than the story. "Head empty, no thoughts" is the perfect way to enjoy It Takes Two.—Moises Taveras
29. Olija
Platforms: PlayStation 4, Switch, Xbox One, PC
With Olija it all comes down to the aesthetic—the muted color palette, the hushed tones when characters speak, the overarching sense of loss and despair that permeates the game. And most notably, those archaic visuals that look like they're from the latest Sierra game you and your friend play on his Tandy computer every afternoon after school. Olija roots its mysteries in the ever-distant, increasingly forgotten past, with all the warmth and sadness that implies.—Garrett Martin
28. Valheim
Driving necessity is at the heart of the survival crafting genre, with games frequently using scarcity as motivation for the player. Valheim abandons this design entirely, instead promoting players to share their resources with one another for the betterment of all. In a genre that so often valorizes individualism, it's refreshing to play a game that both discourages and discredits the practice. Leaning on ideas lent by Terraria, Valheim has created a survival game that has rewritten the rules of the genre for the better.—Nicolas Perez
27. The Great Ace Attorney Chronicles
Platforms: PC, Switch, PlayStation 4
Chronicles is Ace Attorney at its absolute best because it contends with not only its own history as a series but the greater mystery genre that informed its genesis. Moving the point-of-view from a lauded white celebrity to a Japanese man out of his depth is a bold move, and one that shows the glaring flaws in the court system and the inherent racist sentiment that guides it. There's not a single case that feels like a throwaway—each serves as a chapter in Naruhodo's path to understanding his own drive for his profession, and carts the player along a grand adventure that overcomes the somewhat static nature present in the original trilogy. It's an absolute must play for any mystery fan out there.—Austin Jones
26. The Vale
How do you take a "videogame experience" and communicate it to someone who cannot see at all? The Vale has only the most cursory connection to "video" and then almost purely as a kindness to sighted players. The kneejerk response to "a videogame without graphics" is obviously a text adventure, but that would be wrong. While it certainly borrows ideas from text adventures, and video-driven videogames themselves, The Vale has far more in common with radio plays. This is an interactive audio drama. As an audiogame, it delivers an experience in line with big RPG/Adventure titles like Skyrim or The Witcher. And while it might not be the AAA of games for the blind and visually impaired, it might just kick AAA asses into understanding there is both a market for games that cater to these players, and also that there are ways to bake accessibility into existing games that are designed around sighted players.—Dia Lacina
25. Lost in Random
Platforms: PlayStation 5, Xbox Series X|S, Switch, PC, PlayStation 4, Xbox One
Lost in Random is a joy, not just in its shockingly easy-to-grasp amalgamation of gameplay mechanics, but in the entire world Zoink Games has created. Although it lacks the breadth and fidelity of its big budget counterparts, Lost in Random is just as, if not more, immersive and engaging, and it does so within a gameplay system that looks unwieldy but plays like a dream.—Joseph Stanichar
24. New Pokémon Snap
Platform: Switch
Nintendo's relaxing theme park ride of a game tasks you with photographing Pokémon in their natural habitats. Think of it as an on-rails shooter, but with a camera instead of a gun; you'll move slowly along a set path throughout different environments, trying to capture each adorable critter in the most exciting pose possible. Don't worry about taking good photos—just focus on getting each animal in the center of the shot, as large as possible, and ideally either looking directly at you or doing something cute or funny. New Pokémon Snap is a peaceful excursion into a videogame fantasyland full of adorable animals and devoid of almost any stress whatsoever. In other words, it's the perfect game for the late pandemic.—Garrett Martin
23. Super Mario 3D World + Bowser's Fury
No slight intended to Super Mario 3D World, but this game's only on the list because of Bowser's Fury, a short new 3D Mario adventure that's bundled with the rerelease of a Wii U game. Bowser's Fury has one glaring game design decision that will keep it off the list of best Mario games—after a certain point that recurring boss battle sequence becomes an absolute drag—but otherwise it's a fantastically fun 3D platformer that experiments with the classic Mario formula. It almost feels like a rough draft for a future full-scale Mario game, which makes it one of the more intriguing entries in the endless series. Despite being less polished than you'd expect from the usually pristine Mario, though, it's still a wonderfully conceived game that's more than worth playing.—Garrett Martin
22. Mundaun
Platforms: PlayStation 4, Xbox One, Switch, PC
Mundaun's greatest strength is its source material, Swiss folklore. The format, which relies on exploration and puzzle-solving, isn't particularly innovative, but the story it facilitates is cryptic and compelling enough to give it momentum. Its pacing is also wonderfully supported by how well the game blends together its exploration and scripted moments, balancing the two so fluidly that its bizarre events come together in a way that feels almost dreamlike. Its darker moments do not feel cinematically imposed on the player, but rather, that they are something that happens to—or with—them. The visuals, for example, often play on light and shadow in a way that relies on the player's position in the room to progress the scene. Style-wise, its black and white color scheme, often used in similar games to soften rough visual edges (think 2014's Betrayer), combined with hand-sketched textures (reminiscent of Disturbed from back in 2016), evokes the folksiness of a children's storybook but channels a grim sparsity that supports its themes well.—Holly Green
21. Knockout City
Platforms: Switch, Xbox One, Xbox Series X|S, PlayStation 4, PlayStation 5, PC
Multiplayer dodgeball game Knockout City is an absolute blast to pick up and play. It's inexpensive to boot and simple to keep up with, making it markedly less of a chore to log into, have fun with for an hour or two, and hop back out of unlike most service games. It's got a fun style and look to it that makes it all the more inviting, and solid enough mechanics to master that I feel satisfied coming back to practice. Straight up, it's also just fun as hell to play something that isn't so grim or serious, making Knockout City a success in my eyes.—Moises Taveras
20. Black Book
Platforms: PC, Switch, PlayStation 4, Xbox One
At first, Black Book feels familiar. Its card-based battle system borrows from the explosion of deckbuilding roguelikes, most obviously Slay the Spire. The way the game structures itself around gaining new cards and expanding potential strategies will be familiar to anyone who has played games like this before. However, rather than using a slight narrative framing to hold up a number crunching strategy game, Black Book's combat feels like the metaphor of a JRPG. It is a system that deepens its themes of people living in a dying ancient myth. Black Book is interested in a world beyond the material, beyond its mathematical parts. Even as it uses math to represent the ephemeral, it tries to ground the numbers in the mythical.—Grace Benfell
19. Chicory: A Colorful Tale
Platforms: PlayStation 5, PlayStation 4, PC
Chicory is filled with contemplative moments on the purpose of art and how it relates to the mental and spiritual health of the artist. The protagonist soon falls into the dangerous pit of comparing themself to other artists—previous wielders, their sister (a talented artist in school for her craft, while they're just a novice), and others aspiring to become the wielder someday. There's a lot of doubt that comes with being "the chosen one" here, and it's even more salient given the wielder can step down at any time and pass The Brush to whoever they want. It's enough to make the player question their own biases about art; do they value education and skill over enthusiasm? What is the function of art if it's grown to only make us unhappy?—Austin Jones
18. Shin Megami Tensei V
Shin Megami Tensei V's anarchic tone, relatively high difficulty ceiling, and heavy focus on combat might turn away some players, but it has an undeniable gravity and ultimately proves itself as an exceptional RPG. The grind to the top is ascetic and practiced, with grand ambition and keen diligence towards paying the franchise's history its proper dues. It's an undeniably dark game, but quite optimistic in its intentions and eschews edgy clichés. Shin Megami Tensei V is Atlus's contemporary masterpiece, and one of the finest games this year.—Austin Jones
17. Dungeon Encounters
Platforms: Switch, PlayStation 4, PC
Dungeon Encounters might be the greatest dungeon crawler of all time. Bringing together veteran Final Fantasy director Hiroyuki Ito (creator of the Active Time Battle system), rockstar composer Nobuo Uematsu, and Final Fantasy Tactics Advance artist Ryoma Ito, this game is an ATB love letter to Original Dungeons & Dragons. Throw together a party of a Panther-man, a single mom with a medieval shotgun, a golden retriever, even a robot, and go exploring actual graph paper mazes littered with hexadecimal encounters. Dungeon Encounters is the genre stripped to its bones, and made to dance with charm, mirth, and the sharpest approach to combat the genre has ever seen.—Dia Lacina
16. Age of Empires IV
I love inventing guys to be angry with. And Age of Empires IV let's me shout, "YOU TELL EDGAR, I'M FUCKING COMING FOR HIM!" as I highlight a hundred horsemen and press them ever more northward. And each time they bark affirmatively in Old Norman. It rules. When I build new buildings and upgrade my civilization, I can build new guys. Better guys. King William may not have much personality on the battlefield, and largely ceases to exist outside of it, but it's not important. Ages of Empires IV isn't a game about a king's identity. When the Mongols need to differentiate themselves it's through literally picking up their Town Center and moving it around the map—not because The Khan has a real personality to embody. This isn't the game where I'm going to tell the story about the time I, Agamemnon, King of Mycenae, raised an army to chase down one man at the far edge of the world and lay waste to the small township he took final refuge. Initially I thought that was a flaw. Where Total Wars let me become Agamemnon, or Liu Bei of the Perpetual Vibe Check, and Civ V bid me to become Theodora, Hottie Empress of Byzantium, and spread Sapphism across the globe to achieve my glorious Cultural Victory, here there's none of that. I'm simply an amorphous hand guiding the imperial machinery. But honestly, that's okay. There's no distraction in Age of Empires IV. It's a return to a purity of build base, make dudes, obliterate enemies. And I came to love it so much I want to force my dad to spend his weekend playing it with me while he shouts what bits he remembers of the St. Crispin's Day speech over voice chat, as our massive blobs of 15th century dudes collide at our own virtual Agincourt.—Dia Lacina
15. Unsighted
Platforms: Switch, Xbox One, Xbox Series X|S, PC
Would you want to know exactly when you're going to die? Or, worse, exactly when you're going to become undead—when your mind and personality and memories fade away, and you become a brainless killing machine? That's the dilemma at the heart of Unsighted, one of the best written games of the year. A race of synthetic creatures become sentient, driving their human creators to try and destroy them; these automatons now face extinction as the humans deprive them of the element that keeps them animated. Every NPC you meet has a limited shelf life, with an on-screen marker telling you how many hours they have before they run out of juice and become "unsighted." That turns a typical backtracking-heavy Metroid-style adventure into a stressful race against the clock as you try to protect and help as many automatons as you can while also trying to solve the problem of the human threat. What makes Unsighted really shine are your relationships with those other characters; when they start to inevitably blink off and turn unsighted, you'll be overcome with loss and guilt.—Garrett Martin
14. Resident Evil Village
Platforms: PlayStation 4, PlayStation 5, Xbox One, Xbox Series X|S, PC, Stadia
Resident Evil Village goes to great lengths to instill an ominous atmosphere with an odd undercurrent of lightness—there's a ton of dread, like in Resident Evil 7's early moments, but there's an added layer of goofiness that seriously cuts the tension. I love that. Resident Evil has always been goofy; horror games in general are filled to the brim with cheese and insane situations, from UFOs in Silent Hill to dorky dialogue in Until Dawn. Something about allowing the audience to participate in the horror directly through controlling the game's central victim creates hilarious moments, intentional or otherwise. I'll always remember fondly the first time I played Alien: Isolation with a friend and learning the hard way that you aren't actually safe while crawling in a vent. The comedy of horror, derived from inconsistencies in tone and questionable choices no human would make, is an integral element that's simply not acknowledged enough. When I remember a horror movie, I should laugh about my naïve experience sitting through it. I should be eager to terrify my friends with it, to grin as they jump out of their seats.—Austin Jones
13. Hitman 3
Platforms: Xbox One, Xbox Series X|S, Switch, PlayStation 4, PlayStation 5, PC, Stadia
Playing Hitman 3 feels like being thrown into a random improv scene. You're constantly switching parts, objectives, and wardrobe, making sure never to break character before you eliminate your target. Every stage is a performance, and they're all incredibly distinct and fun. You play as world-class executioner, Agent 47, and due to his occupation there's a thick cloud of intensity and death that follows him around every corner. Each contract sends you to varied and visually striking locations all over the globe, setting up flexible environment-specific boundaries while simultaneously encouraging you to push against them (or even throw it all out and do it your way). Both the plot and general premise of IO Interactive's Hitman 3 are straight-faced and sober, yet it somehow manages to be one of the funniest games I've played in a minute due to clever prop comedy and witty, well-written NPCs. After spending a bunch of time playing around in its charming and compact world, Hitman 3 has proven to be a well-constructed assassination sandbox full of tension, fashion, and possibility. —Funké Joseph
12. Before Your Eyes
Don't dismiss Before Your Eyes as a gimmick. Yes, it's the blinking game—the one that you literally play by blinking your eyes in front of a webcam—but there's way more going on here than this mechanical tomfoolery. With its focus on a newly dead man's entry into the afterlife, it has a touch of Pixar's Soul crossed with the mundane sorrow of something like Death of a Salesman. The game looks back on the main character's very normal life and the major relationships that shaped it, and the story's generally well-told enough to avoid the kind of cheap, maudlin sentimentality you might expect from it. The blinking contributes more to the game's power than you might expect, but if you don't want to hook your webcam up you can always play it with a mouse instead. Before Your Eyes is a small, quiet game with outsized emotional heft.—Garrett Martin
11. Deathloop
Platforms: PlayStation 5, PC
At the end of the day, the loop is all that matters. Deathloop leaves it all out on the table, reconstructing the immersive sim a few different ways in order to pull you under. Luckily for the game, it's a damn marvel and works like a charm. While I can nitpick about Deathloop's shortcomings, I'd rather just point you to a game that's a joy to play, confident in itself, touts two wonderful Black leads, looks wonderful, and rewards you for thinking outside the box. While it doesn' quite feel like an evolution of the formula, it's almost assuredly Arkane's most feature-complete and refined take on it. Deathloop is countless things, and most of them are great.—Moises Taveras
10. Loop Hero
Loop Hero is a roguelike, a deck builder and an RPG with the cadence and look of a tower defense game, wrapped in a grim but simplistic '90s PC game aesthetic. It's a mashup that feels like it shouldn't work, because that sentence I just wrote sounds preposterous. Instead, Loop Hero is absolutely magnificent. While it may seem unengaging because it effectively plays itself, it really is just prompting the player to look at gameplay from another angle, namely a more systems-driven one. For a person like me, who doesn't really craft "builds" in RPGs, it's made me realize why that is actually a rewarding aspect of those games. Now I spend half my time in Loop Hero making numbers go up and making optimizations I never would have, before embarking on another loop.—Moises Taveras
9. Genesis Noir
Platforms: Xbox One, Switch, PC
Genesis Noir is a cosmic point-and-click mystery about the meaning of life, the tragedy of death, the creation of the universe, and, oh yeah, jazz. Yes, it's incredibly pretentious, but in a way that absolutely works, drawing you in instead of pushing you away. It has lofty goals and it isn't afraid to really go for 'em, with a cleverness and thoughtfulness that makes even the most esoteric decision land with power. It's also the most stylish game of the year, with a noir-ish black-and-white color scheme occasionally broken up by flashes of color, incredible character designs, and an atmospheric jazz score that fits it perfectly. It's one of the most beautiful and entrancing games of the year.—Garrett Martin
8. Monster Hunter Rise
Monster Hunter World almost got there, but Monster Hunter Rise feels like the first game in Capcom's smash series to offer something substantial beyond the repetition of monster hunting. There's a real world here, with fleshed-out characters, and it makes Rise a pleasure to visit even when you aren't looking to slay some beasts. And when you do take up the hunt, the clearly regimented quest system makes it fit perfectly within your busy schedule—you can pick up the controller and know you'll be able to knock out a mission in under a half-hour. It's a role-playing game that doesn't demand all of your time, which is exactly the kind of RPG we need these days.—Garrett Martin
7. Wildermyth
Decisions in Wildermyth aren't about getting something right or wrong. They're about telling the most entertaining story. And that's why no videogame has ever been more reminiscent of my tabletop RPG experiences with friends. Rather than a system like Dungeons & Dragons that offers small variants of failure and success in actions, it is more similar to modern systems like Powered by the Apocalypse games and Blades in the Dark, games that are designed for players to "fail forward." It's a thematic value that I wish more games implemented.—Waverly
6. Forza Horizon 5
Platforms: Xbox Series X|S, Xbox One, PC
Forza Horizon 5 is the most gorgeous and dynamic game I've played on the Xbox Series X by a mile. That said, the beauty of the game isn't just in the mechanics alone. It's in how the game loves, respects, and brings Mexico to life. When it comes to representing Mexico on-screen, Americans like one thing, and pretty much one thing only: Sepia tones. Baked in browns and oranges, representations of Mexico on screen in film and television strip the country of its beauty and distill it down into its most stereotypical parts, often using it to highlight narcos. But in Forza Horizon 5 the diversity of the races is met with the diversity of Mexico itself. Mexico isn't just a desert landscape, and the 11 distinct biomes in the game highlight that. It's clear that a lot of love went into Forza Horizon 5. You can see it in the car selection. You can see it in the environmental design. You can hear it in the playlists. This game thrives on a culture of love that is baked into every gameplay element. In every way, Forza Horizon 5 is a love letter to Mexicans, and it's one I'm thrilled that I opened.—Kate Sánchez
5. Sable
Platforms: PC, Xbox Series X|S, Xbox One
Fittingly, Sable's title comes from its protagonist's name, not her role or the place she lives. It foreshadows the game's small stakes. Sable is a Glider. Just now coming of age, she leaves her home to discover who she might become and what role she might play. To begin, Sable wears the mask of her culture, made from an Ibex skull. As Sable helps other people, she gets badges, which she can turn in for other masks. For example, she could take the mask of the mechanist becoming an expert on refashioning the ancient machinery around her into something practical. She could become a Climber, exploring the highest margins of the world. She could become something as yet unnamed or unanticipated. Its openness means that Sable makes very few demands of you. In a real sense, everything is optional. Because of that, it actually feels free. It is not the freedom to move through space uninhibited, to dominate or control. Rather it is the freedom to determine who you are, to let the people around you make you into something new.—Grace Benfell
4. Death's Door
Platforms: PC, Xbox Series X|S, Switch, PlayStation 5, Xbox One, PlayStation 4
Death's Door implicitly argues something the entertainment world at large needs to understand: Nostalgia doesn't have to be shameless or oppressive. It doesn't have to be the summation of a game's (or a movie's, or a TV show's) ambition. It doesn't have to be splashed all over the cover and title screen, or the full extent of the marketing campaign. Death's Door deeply evokes the spirit of some of the most beloved games of all time, and does it well enough that anybody familiar with those legendary games will no doubt recognize and appreciate it. And it does all this with a context and presentation that makes it feel new and vital and not just like a calculated imitation of the past. It takes so much of what made the original Zelda and A Link to the Past into timeless classics, but makes them into their own. Nostalgia can be part of the package, but it shouldn't be the whole point, and Death's Door's cocktail of mechanical nostalgia and narrative creativity is something we don't see enough of in today's IP-crazed business.—Garrett Martin
3. Metroid Dread
The first entirely new side-scrolling Metroid game in almost two decades captures what makes Super Metroid such a timeless classic, while also introducing something the series hasn't seen since the very first NES game: a palpable sense of fear. The almost unbeatable E.M.M.I. sentries are even more terrifying than the metroids you face at the end of Metroid. Every E.M.M.I. encounter punctuates the game's sterling design with genuine dread, eliciting an emotional and physical response rarely seen in games. Metroid Dread is about as good a return to classic Metroid as anybody could ask for.—Garrett Martin
2. Psychonauts 2
Platforms: Xbox Series X|S, PC, Xbox One, PlayStation 4
Psychonauts 2 feels like a game made by real people who care about real people. Many games have come down the pike the last several years with a focus on the psychological state of its characters, and thus its players, but too often they lack any tact or any legitimate insight into how people think and feel. They use sorrow and violence as shortcuts, relying on cheap scares and easy provocation. It's like they're made by machines, or the board room, or some algorithm that slightly rearranges previous AAA hits into something that's supposedly new. Too many of these games fall into that witless trap of thinking something "serious" and "important" must also be humorless and dark, unrelentingly grim and fatalistic. Psychonauts 2 reveals that for the nonsense that it is, showing that you can more powerfully and realistically depict emotion when you use warmth, humor, humanity—the whole scope of emotions that make us who we are. Psychonauts 2 asks "how does it feel to feel?", and then shows the answer to us—and the games industry at large—in brilliant colors.—Garrett Martin
1. Ratchet & Clank: Rift Apart
The first real reason to own a PlayStation 5, Rift Apart is an embellishment on a formula that's worked for 19 years. It's splashy and charming. It doesn't take itself too seriously, but it absolutely will dive into talking about trauma and disability, and tackle questions of belonging and imposter syndrome in ways that are simple enough to speak to children, but honest enough to resonate with adults. And at times, it manages to be surprisingly funny despite being entirely predictable, knitting trope to trope in a tapestry wrapped in more tropes. It's a simple but surprisingly earnest and compassionate game. What carries this big flashy sci-fi romp along and helps elevate it from a simple farce is this charm and humanity. Rift Apart has the heart that Guardians of the Galaxy could never find.—Dia Lacina
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Q: magento 2.4 category_form.xml add comment to fields I'm creating some custom fields for my categories, so I've added a couple of simple text fields in the database with the setup php script, then extended the category_form.xml to show my fields in the admin category page. Everything works fine: I see the fields in the admin category page and I can save the values.
I'd like to add a line of comments under those fields. I thought I could just use the comment tag like I do in every other xml form but here I get the error "Element 'comment' is not expected".
Is there a way to add a comment-like sentence under the field?

A: there is a note field in the eav_attribute table. This field is used to store the label that appears below the fields in the admin screen.
As a consequence, you can add a data patch that updates the category attribute you have created so that the note's value for this attribute has the label you want to set.
Below is an example for product at \Magento\Catalog\Setup\Patch\Data\UpdateProductMetaDescription. I did try manually for category and it works indeed.
/** @var EavSetup $eavSetup */
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            ]
        );

redpajama_set_name: RedPajamaStackExchange


Hi, I'm Jennifer Maggiore with redballooninc.com, and I'm going to explain how to switch from auxiliary mode to video mode on your TV set. You'll simply get your remote control and look for a button labeled input. It might also be labeled TV/Aux or TV/Video. You'll then continue to click that button until you go through all of the different options which might include some other options such as compositive or HDMI, and you'll just continue to cycle through those options until you reach the one that you're looking for. That's how you switch from auxiliary mode to video mode on your TV set. I'm Jennifer Maggiore with redballooninc.com.


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redpajama_set_name: RedPajamaGithub


Day 1: Friday, January 9, 2015.
11:00ish: Scheiter et al. Modeling functional trait diversity. aDGVM (adaptive Digital Global Vegetation Model) does very well simulating African savannas. Does it do well with Australian ones? Not at any useful scale, so they re-parameterized for Australia. This means that e.g. stem diameter doesn't relate to canopy diameter & height the same way in these two systems. I would suspect phylogenetic constraint….
Today, many models use PFTs – plant functional types – to parameterize; lump species by shared functional traits to keep system simple. aDGVM2 is a trait-based dynamic veg model, uses an individual-based approach, so it doesn't lump species by PFTs. Models natural selection, mutation and cross-over to optimize for productivity. Cool. The model produces emergent coexisting life history strategies: these are not programmed in, but arise from the interaction of the digital plants and the physical environmental parameters.
11:23: Ruiz-Benito et al. Trying to understand how 21st century climate change will affect functional diversity of European floras. The current consensus seems to be that Europe will lose tree diversity and overall biomass, producing a carbon source. These authors want to know whether higher functional diversity might protect some forests from climate change. FD had an effect on growth and regeneration, but not mortality, but Functional Identity had an effect on mortality. These effects were different in evergreen, broadleaf, and Mediterranean forests, so those cannot be expected to respond to climate change in parallel ways.
11:35: Hawkins et al. Begins with an apology: there will be no biology in this talk. I do a silent happy dance. Also, he says that his papers are sets of maps with filler text, which seems a good idea to me. Now, onto the science: begins with a division of drivers of community structure into extrinsic and intrinsic variables. Traits are intrinsic variables. He now suggests intrinsic drivers may not even show a correlated pattern, let alone mistaking correlation for causation. I'm not sure I understand this point. He shows that cherry-picking random 'traits' can produce patterns that fit any particular hypothesis. I think the point of this talk is that dumping trait data into biogeographic models (of, say, richness) you can get very high R-squared: many of these methods seem to overfit as a consequence of the spatial patterning of the species. Species aren't randomly distributed, so models can see patterns where there are none. This problem also appears when modeling random traits to predict random macroecological values (modeling, e.g. Functional Diversity). An issue I see here: the apparent strength of random predictors means that even if you're not statistically fishing, but testing a hypothesis, you cannot be confident that you're actually seeing a biologically meaningful effect. What's the answer? We don't know yet, but we're a smart community of scientists and we can work on this.
The jet lag got me really bad for the rest of Day 1, so I spent the time until the poster sessions chatting with colleagues in the hallway and procrastinating working on grant proposal and classwork.
This is the paleobiogeography session! I'm in my element.
8:30: Stigall gives an introduction and overview of the principles of paleontology for the neontologists in the audience. I'm a jetlagged zombie, so I'm zoned out through most of it. She has a critical part at the end where she distinguishes between completeness and adequacy in the fossil record, beginning with Darwin's apologia for the incomplete fossil record and then reviewing 1) Phylogenetic completeness, 2) Stratigraphic completeness, and 3) Geographic coverage. As it happens, the fossil record is known adequately in these three ways to test many, many paleobiogeographic hypotheses.
8:52: Servais and Harper on the origin of biogeography. They are from a different world, not digging in Quaternary mud, but doing real, hard fieldwork in real rocks… Mentions Wegener and Continental Drift, but Servais and Harper are working before Pangea. How can they reconstruct the shape of Earth's geography before Pangea? Let's find out… In the end, you have to pull together all of the lines of evidence, from structural geology, sedimentology, etc. etc. (Note: Pangea looks an awful lot like Pac-Man. Coincidence?). Then you make a hypothesis and test it with new data. Science!
Now he's linking Sepkoski's paleodiversity curve to break up and unification of supercontinents: Times of lower diversity have been supercontinent times (Rodinia and Pangea) and times of higher diversity have had distributed continents (like now!). Servais et al. (2009) GSA Today.
This example is followed by several others that I won't record because I'm feeling lazy.
9:15 Kiessling and Kocsis: Marine benthos over last 300 million years… Starting out with Valentine's seminal 1970 work on species richness over geologic time. He explained this pattern with an increase in provincialism after K-T. However, Miller et al. 2009 used the PaleoDB to show no significant change in provincialism through the paleo record.
Kiessling points out that the PaleoBioDB is millions of records freely available to everyone. It's a great resource.
Then a little dig at Servais' talk, pointing out that you cannot use paleocontinental reconstructions from before Pangea because the researchers are still moving the continents around. So, he's focusing on the Pangea to today part of the record.
Climate context: Warming and cooling in two cycles from Permian to today.
He's fine-tuned a system to diagnose biogeographic provinces in marine inverts over time, producing a movie of province changes through time. He has good evidence that the number of provinces is driven strongly by sample size, so must standardize for sampling. Once you do, the variation in number of provinces through time disappears. The only interval that seems to stand out is a dramatic loss of provincialism in the wake of the P-Tr extinction.
9:39: Denk, on whether or not we need fossils for historical biogeography. Shifting to Cenozoic terrestrial plants. Then he uses quotes to show that over the last 20 years, biogeographers have shifted from shunning fossils as unnecessary in a Bayesian phylogeographic context to part of a continuous paleo- and neo-biological dataset. I can dig it.
Example 1) Smilax, greenbriar. Reminds me of Smilex gas from the old Tim Burton Batman movie. Good fossil record, good phylogeny. Four clades identified in recent phylogenies. Three new-old world disjunctions in these four clades. Mid-Miocene fossils of Smilax in Iceland suggest the disjunction of some clades might be because of a bird-accessible North Atlantic crossing. Maybe not quite a continuous land bridge, but a convenient bird corridor. Plus, fossils show that the one clade that currently has no new-old world disjunction did, back in the Miocene. Really, everything was just better in the Miocene.
Example 2) Quercus, the oaks. They have a rich fossil record and a strong, well supported (almost oak-like?) evolutionary tree. See what I did there? The surface texture of the pollen grains is distinct among the deepest-diverging branches of the oak tree. Heh. Oak tree. The white oak tree seems to germinate in the high arctic of the Eocene and then spreads its leafy boughs over the temperate zone with climate cooling through the Tertiary.
Example 3) Dracaena, dragon trees. There's no phylogeny for this group: sad because there's no Dragon tree tree. Modern distribution: disjunct distribution on coast of western, eastern, and southern Africa. Middle Miocene fossils from Turkey! Fossils suggest the ancestors of modern disjunct species had distinctly different ecology from modern groups. Convergent ecological divergence among these lineages? If you had a tea tray with the phylogeny of Dracaena on it, it would be a dragon tree tree tray. And.. if someone was giving away carts made from Dracaena, you could get a dragon tree wagon free. If someone carved the phylogeny of dragons onto one of these trees, it would be a dragon tree dragon tree.
10:33 Sanchez Meseguer et al. The present isn't always the key to the past, when there is sampling bias (e.g. extinctions in phylogenetic trees of extant taxa). This problem can have a major effect on reconstruction of ancestral areas. For a moment, clouds part and sunlight obscures the screen, but then the gray day returns and the lecture continues.
Hypericum example. Originates in the palearctic and doesn't spread to the rest of the world until the Pliocene. Modern phylogeny says that it originated in Africa, Europe, and North America with previous clades in Africa – disagreement! So, they added ancestral inferences from fossils. Next, they fit niche models to fossil distributions and paleoclimate simulations to get climate tolerances for the earliest members of the genus. In this way, they could see climate barriers to distribution in Beringia and equatorial regions. These climate data explain the palearctic limitation of the clade until Pliocene cooling. More detailed analysis also suggests bias in extinction with more eastern palearctic lineages going extinct, producing the erroneous pattern in the phylogenetic reconstruction of ancestral regions.
10:48 Badgley – Continental gateways and mammalian faunal change. Four-part biogeographic states: permeable/impermeable barriers and changing/stable climate. Four unique macroecological predictions for the four states.
Example 1) Siwaliks of Pakistan. Focus on mammals >1kg. Floral turnover from C3 to C4 grasslands through this sequence (told from carbon isotopes). Most of the sequence has endemic species in genera shared with other Asian sites: so regionally similar on the broad scale, but different in details. Three of the four biogeographic states recognized in this sequence. 14 out of 16 predictions upheld by observations, suggesting support to this model of climate/barrier interaction.
Example 2) Miocene mammals of Iberian Peninsula. Three analysis intervals, with two of the four states. Environments reconstructed from paleobotanical remains + stable isotopes from mammal teeth. Domingo et al. 2014 Paleobiology. 10/12 predictions upheld by evidence.
Example 3) Quaternary mammals of Cape Fold Belt of South Africa. (Faith and Behrensmeyer 2013, Paleobiology). Area of ecosystems and connections fluctuate with sea level change during glacial/interglacial cycling. At Pleistocene->Holocene, ¾ predictions met, and the last one, too, but in a nuanced way. In particular: grazers hardest-hit by the climate change.
He's asking us all to visualize a heatmap of global species richness of our favorite group… Then he goes to the Fine et al. 2006 (book chapter: no link) plot of tree richness and unlogs the diversity axis: the diversity of non-tropics is like a thin green veneer compared to the diversity of the tropics.
Now squamate reptile richness: snake richness peaks in the tropics. The other 'lizards' add a peak in Australia (in desert scrubland: a desert 'coral reef', he explains). Other environmentally comparable deserts have nothing like this diversity of lizards. An example of a non-latitudinal diversity anomaly.
What processes can cause these gradients? Mittelbach et al. (2007) Ecology Letters framework: Ternary diagram of Ecological/equilibrium vs. diversification rate vs. time.
Today's focus: "Speciation rate" is not a single hypothesis.
Rhode (1992) Oikos: higher metabolic rates in the tropics.
Biogeographic species pumps: Mosaic habitats of Australian arid zone. Pianka (1972) Copeia: Habitat fluctuations lead to speciation.
These all predict that speciation rates should be higher in hotspots. Does speciation rate explain major gradients in vertebrate diversity? (I like that he has an actual question with a '?' here).
Now an example from Australian sphenomorphine skinks. Most diverse terrestrial vertebrate radiation in Australia. Big, big tree. Rabosky et al. (2014) Sys. Bio. Creates Bayesian 95% credible set of rate shift configurations.
What about extinction? He's not going to interpret extinction rates from molecular phylogenies. One would need to integrate fossils to get any kind of confidence. Davis (no relation to me!) et al. (2013) BMC EvoBio. Testing effectiveness of reconstructing speciation and extinction rates from model phylogenies. Any phylogenetic tree is compatible with a range of extinction events from zero to infinity.
Nee et al. (1994) Phil Trans B: phylogenies contain max recoverable information about species rates towards the present.
Example: Speciation and geography of marine fish diversity. Collaboration with a bunch of people, whose names I didn't get because I'm a fail slow typer.
Is the Indo-Pacific a species pump for fishes? BAMM analysis across whole living fish tree, translated to the map using their spatial database. Then a map of mean speciation rate for all available grid cells. Surprise: the highest rates are in the Arctic and Antarctic! It's like the speciation rate is the inverse of the latitudinal richness gradient. So, instead take the latitude-midpoint of each species, but it still produces a smile-shaped plot with highest rates at the high latitudes. Then he goes through a bunch of specific examples: Arctic/Antarctic clades have high rates of speciation. Famously diverse tropical fishes have low speciation rates.
What's driving this pattern? "I have no idea." But he can tell us what it isn't.
Short on time, so one more story in 4 quick slides. Whew! This talk is a wringer.
NW land birds dominated by tropical sub-Oscine clade. Now plot rates on using BAMM. Lots of clade-specific variation in speciation, but no relationship with latitude. Tropical and temperate rate distributions are identical. See also Jets et al. 2012 and Kennedy et al. 2014.
We can make high-res macroevolutionary rates.
Need the fossils to do extinction, and he's extended BAMM to work with total evidence trees that include fossils (yay!).
No matching of speciation rate with high species richness. Evidence against a broad class of mechanisms that have been invoked for creating species diversity hotspots. Not a lot of evidence for species pumps, at least for vertebrates. Plants, maybe different.
Lunch was good, chatting with Catherine Badgley and Kaitlin Maguire about neoendemism projects.
I had talks picked out for after lunch, but I'm really feeling the jet lag now, so I'm taking a mental health break instead. Looks like the next talk for me is at 14:30. I'm really looking forward to the evening's activity: banquet at the steam train museum!
2:48 Renner volunteers to not give her talk to get the session back on schedule, but there is a general outcry from the audience, so she goes ahead! So, how to assign fossils with too few characters to place them precisely on a phylogeny? Don't want to throw out those data. My battery is dying, but this talk is derived from a 2014 paper, so you can check it out later. Heath et al (2014) PNAS.
8:38: Rodrigues-Sanchez introduces the session on using past climate to inform predictions of response to future climate change. Cites Clark et al. (2001) Science: Need a better ability to make ecological forecasts. Emphasizes 1) climate is changing and we need to anticipate the future and 2) climate has always been changing, so we can use past data to test models of ecological responses.
What can we really learn from the past? What? E.g. species response to past climate variations. How likely? How often have species had consistent responses to climate change? Why? What processes are actually controlling range change, plastic response, and extinction?
8:47 Steve Jackson, begins by pointing out that the last IBS he went to was the first, and his talk there was his last one using 35mm slides! Times have changed quickly.
Now he's going to give some for-instances of how paleoecology can help conservation biogeography.
How sensitive are natural populations and communities to climate change?
How are they able to persist?
Long-term population persistence – example populations >10k yr in one place.
What are the biodiversity costs of climate change?
Examples, including Tony's Irish Elk study.
How and how far do species move in response to climate change?
Note from me: He's using 'migration' for this movement, but I would not.
Migration is a regular, cyclic range change as part of a species' natural history.
This sort of range change might be called immigration or emigration, but it is one-time response to climate change over multiple generations, so not migration.
How to discriminate between real 'migration' and apparent 'migration'.
Example Williams et al. 2004 Ecol. Monogr. Fagus grandifolia moves north, but McLachlan et al 2005 Ecology DNA shows that they were already in the north.
What are the relative roles of environment, biotic interactions, and chance on ecological and biogeography outcomes of climate change?
Figure from Jackson and Blois 2015 PNAS (in press). Extent by duration in log-log space with processes shown at different scales. I need to find this figure.
Need to carefully consider the complex interactions among these factors.
Wraps up with a call for better interdisciplinary interaction, better consideration of taphonomy, and overall epistemological rigor. And don't forget natural history: these are real individuals in real species, real ecosystems.
Know what's at stake for decision makers: what are the consequences of errors? Scientists have relatively low consequences for being wrong. Not so for conservation planners.
9:13: Morlon on past effects of climate changes on biodiversity.
Current geographic gradients suggest climate has a control on diversity, morphology. Similarly when you look at species richness/traits versus temperature in the fossil record (example from Ceonozoic pollen, brachiopods). Gilooly et al.2001 Science: temperature affects metabolic rates, body size.
So, she presents some hypotheses about the processes that would link temperature changes on speciation rates. Overall hypothesis is that speciation rates should be positively correlated with temperature. Similarly, extinction rates should be negatively correlated with temperature, and rates of phenotypic evolution should be positively correlated with temperature.
How to test? Traditionally, using modern gradients and the fossil record. E.g. Ezard et al 2011 Science. However, looking only at the modern ignores the contingent evolutionary history. Now she just suggests that some groups (birds, insects?) have fossil records too poor to be used for this type of study. That seems too defeatist to me (referring back to Stigall's adequacy vs. completeness bit from yesterday).
Now she's onto the idea of using genetically-based phylogeny to make tests of these hypotheses using comparative methods. E.g. Nee et al. PTRSB 1994. Morlon et al. 2011 PNAS, Morlon et al. 2014 ELE.
Condamine et al. 2013 ELE. –> Incorporate external knowledge of temperature, sea level change, etc. into modeling of rates on phylogeny. Interesting. New in prep paper tests 21 different models of the way temperature affects rates. Results for mammals: support positive correlation between temperature and diversification rates, as expected from hypothesis.
Cantalapiedra et al 2013 PRSB: similar results for ruminants.
So: could decreasing temperatures in the Cenozoic explain the widespread observation of declining speciation rates over time through many comparative studies?
One of her collaborators is working on incorporating environmental dependence in Brownian model of trait evolution. Interesting: signs point to an inverse relationship between size evolution and temperature (faster trait evolution in cold intervals). C.f. Cooper and Purvis 2010 Am Nat.
She hopes to convince us that phylogenies can be useful for understanding the interaction between evolution and climate change. Quest completed.
The Q&A focuses on two things: the disconnection between speciation rate and mass evolution rate and the problem with using global climate to model a local process (natural selection/speciation).
How natural populations respond to environmental change: move, go extinct, or evolve.
Need to understand feedbacks between phenotype, environment, and genotype. Problems: phenotype-genotype link is only established in model species, but with little knowledge of their link to environment. Similarly, ecological model species are not known in genotype-phenotype linkage. She would like to close this loop.
Need accurate measurements of effects of environmental change on natural populations. But we have no long-enough-term datasets. How can we do this without investing multiple human careers? Using omics on short-generation time species in the lab, like yeast. Also, we can use space for time, e.g. Gillespie et al 2013.
But here is her new proposal: Orsini et al 2013 TREE. Evolutionary time machine: sample at different spatial scales with multiple time series. It looks like she's integrating across multiple sediment cores taken from a single lake. Yes, she is looking at waterflea microevolution over space and time. Aha! She can re-activate resting stage waterfleas trapped in the sediment to see what different time steps do in "common garden" experiments. She has several citations here: Orsini et al 2011, 2012, Pfrender et al. 2012, and others. This seems a very elegant use of natural experiments, isolating sediment cores that have one or another factor controlled so she can see what each one does to the local lineage. They found a clear genomic signature of natural selection and repeatable patterns of local adaptation.
This project shows the prospect of a new field of paleo "omics".
Now for the coffee break, so I'm off to put my presentation on the computer. Except… There were no people or computers in the room, so I'll get to put my talk on the computer at the last minute. As one does.
10:37, Fordham: Extinction risk from anthropogenic change, fossils and molecular log books.
Moving beyond predictions: using predictions to improve reserve design, plan translocations.
Wants to use the late Quaternary as a natural experiment to improve predictive models for policy makers. Again, emphasizing the need for integrated approaches across paleo-, neo-, et al. This is a refrain of the morning's presentations: if we want to actually save any bits of biodiversity or natural ecosystem function, we need to all work together.
By combining aDNA, climate, and conventional paleoecology data, one can create a model of the demographic changes that led to extinctions in the past, providing an empirical basis for extinction risk given current or expected demographic shifts.
Is our current high forecast biodiversity loss a real thing or a consequence of model bias? He makes a good point here: testing climate biodiversity models using 20th century data can't capture the range of expected climate change into the 22nd century. You MUST go back to the Quaternary fossil record.
OK, back to the talk: Mechanistic underpinnings of climatic refugia. Blois et al. 2011 Quat Sci Rev. Williams et al. 2000 Journal of Biogeography.
11:05, Jansson et al. Effects of stability on bird diversification.
Dynesius & Jansson 2014 Evolution: does change or stability foster diversification more? Reminds me of the old Plus ca change model of Sheldon 1996 Palaeo3.
74 sister groups of birds from New World. Used niche models to calculate areas in LGM and today to measure persistence. They're using genera and one of the characters seems to be species richness within the genus. They're using sister genera so they can do PIC on the pairs.
Climatic stability was significant to diversification only for tropical genera. Interesting.
11:21, de Boer and Hooghiemstra: Paleoecology of Mauritius. Settled by humans very late (1600s), so excellent natural lab to distinguish human from natural ecological perturbations.
Cool: mass-death assemblage records 100 years of dying during global megadrought 4200 years ago. Sediment cores indicate the blooming of toxic cyanobacteria during this interval, killing the vertebrates using the waterhole.
Pollen record from the central crater shows stability until the start of the Holocene, at which time there is a revolution in the island's ecosystem.
Lowlands and uplands have different disturbance regimes, do not respond the same. Take home: be sure you have analogous systems before extrapolating from one to another.
The Q&A seems centered on whether or not he can call his bone assemblage built up over 200 years a 'mass death'. In the end, it is a terminological concern, but I'm not sure the two sides are understanding one another.
aDNA to get at demographics through time.
Models based upon these data to forecast range dynamics using current genetic diversity data.
Modeling niche space through time, instead of as a static thing from modern time slice.
Another call for integrating data and methods across disciplines! And he wrapped fast to keep us on time.
I led the paleobiogeography student discussion table at lunch. We had a lively chat, ranging over topics from the climate zones of Oregon to how to use niche modeling techniques to predict fossil localities.
1:01, Hoogiemstra et al.: Islands and sky islands.
Sky islands in the N Andes. Most of these islands have existed in this way for <5Ma. Defining sky islands by their tree lines. How much time is available for connectivity among these islands? There is a lot of fluctuation in the tree line, with the highest expression only 2000 years ago. The modern high tree line and divvied-up sky islands are the exception, only about 10% of the time in the Pleistocene. You must test ecological theory against glacial settings, more like 60% of the time. He's suggesting that these sky islands are not obeying the MacArthur and Wilson island rules, because they have not lost species richness, despite losing 96% of their expressed area since the glacial maximum.
Now ocean islands. Where is the species pool for the change at the onset of the Holocene in Mauritius (see earlier talk)? He suggests the gallery forest. Also, the forest is down to 2% of pre-human levels on the island, but they still have high species richness… The unanswered question: how long can it last?
1:33 Gutierrez-Garcia et al. Opens with a Jurassic Park reference: when it came out she thought there was little chance of getting any DNA from any fossil. No longer. However, there are no current tropical aDNA sites. Loltun cave: Late Pleistocene to Holocene fossils. Ototylomys phyllotiys, an extant central american mouse, is found in the cave. Nice! They made a custom 3D scanning system just to get a surface model of the tiny, tiny jaw they were going to destroy for aDNA. Wow! The cave sample made a new, basal clade outside the entire crown group of the species. (Gutierrez-Garcia et al. Biology Letters vol 10).
Q&A points out that they could not micro-CT the specimens because it would cook the aDNA. Hence the super fancy structure-from-motion micro scanner.
Trophic downgrading and loss of megafauna is a big deal today, and there is a lot of public support for protecting the big animals. Cites Estes et al. 2011 Science paper about trophic downgrading in the wake of large animal loss. Felisa suggests we can use the wake of the paleoextinction to model what to expect in our neo-extinction future. She's going to tell us what happened to the survivors in the Hall's Cave, Texas, fossil record. Pre-extincntion, the cave record has a very large number of species, many of which are megafauna. Only 3 of the megafauna species survive. The body size distribution changes in the wake of the extinction, and not just in the loss of megafauna. There's also a bigger turnover in species composition than the change in mass distribution. Details of PAIRS results wash over me in rapid succession. Take home: more significant segregations than associations. Many pairs of species were pushing one another out.
2:03, Milan Chytry et al. A refugium in southern Siberian mountains.
European full-glacial fossil sites often record tundra, taiga, and steppe organisms together for both plants and animals. A no-analog community? He asks whether there is a potential modern analog. Mammal and plant distributions of tundra, taiga, and steppe organisms coincide in southern Siberia. The pollen record confirms the hypothesis that this area matches the unusual full-glacial sites. The snails also agree. So do the mammals.
2:18, Willner et al. Florsitic legacies of Quaternary climate change in European Beech Forest.
He's looking at the distribution of the understory plants from beech forests to see if they have a pattern of distributions that matches the hypothesized refugia. In the end, they do.
I skipped the next session of talks to chat with folks and be generally zoned out.
Now up: the Wallace Award, awarded to Simberloff this year.
Character displacement and release in the small Indian mongoose and the stone marten.
Links his topic to Wallace through Wallace's work on the effect of introduced predatory mammals on islands.
1872 introduction of Mongoose to Jamaica –> there to kill rats, it became one of the worst 10 invasive species. It was subsequently introduced to many islands worldwide, and it has always been a disaster. In its native range, it's the smallest of three mongooses, but in all introduced areas, it was put into a carnivore-free system. So: it should make a good test of Brown & Wilson's 1956 ideas about character displacement and release. Would it get bigger in the absence of competition?
Measured two traits: skull length as proxy for body size and upper canine length.
Simberloff et al. 2000: first publication on this project. In most islands, males got bigger and females did not, so sexual dimporhism also went up.
Mediterranean islands off Croatia some have stone martens as well… Hypothesized to defeat male body size increase. Yep it seems to work.
The problem is in the exceptions: Several islands do not fit the trends.
Now details about Stone Martin distribution. Interestingly, the invasive island forms are smaller.
The mongoose won't cross water without help, so rivers are important boundaries for them.
The data are broadly consistent with the hypothesis of character release.
So ends my notes of #IBS2015. I look forward to the next meeting, January 2017 in Brazil!
BAMM does not involve any sort of "machine learning".
« In which we explain how camel ankle bones relate to the fate of global ecosystems.


text:
Dragon with an Agenda
A Dragon with an Agenda is a character serving as The Dragon to the Big Bad, but having different goals from him. For example, if this character encounters the heroes immediately after the death or defeat of his boss, he won't try to complete his master's Evil Plan, but will instead go on to pursue his own plans.
Most commonly, he and the Big Bad have a mutually profitable alliance, and the Big Bad just happens to be more powerful of the two. If he's also influencing the Big Bad's plans, then he's at the same time the man in front of the man and The Man Behind the Man. Dragon-in-Chief is when The Dragon is actually the more dangerous of the pair, by a significant margin. It is not unknown for him to have taken on service for his own purposes and fooled the Big Bad into thinking he's subordinate.
Rather than a straight evil character, he tends to be an Anti-Villain or an Enigmatic Minion. Contrast with Battle Butler and Psycho Supporter, who are often The Dragon but have the same goal as their master. Unlike The Starscream or the Reliable Traitor, they are usually at least nominally loyal, and their main agenda doesn't outright conflict with that of their boss. If The Dragon and the Big Bad are equal or nearly so, you have a Big Bad Duumvirate. If they outlive the original Big Bad, this type of Dragon may go on to pursue their own motivations and become a Dragon Ascendant. Compare/contrast with the similar trope Piggybacking on Hitler. The equivalent among the hero's allies is ...Who Needs Enemies?.
Examples of Dragon with an Agenda include:
1 Anime and Manga
3 Fanfic
4 Films -- Animated
5 Films -- Live Action
7 Live-Action TV
8 Tabletop Games
9 Video Games
10 Web Comics
11 Web Original
12 Western Animation
Anime and Manga[edit | hide | hide all]
Archduke Gorgon from Mazinger Z and Great Mazinger. Theoretically he allied himself with Big Bad Dr. Hell to help him to destroy Mazinger-Z. But his real goal was watching over Hell and betraying him at the right time to get him killed (in the GosakuOta manga he literally backstabbed him) and Mazinger-Z destroyed. He succeeded on all fronts.
Scary Black Man Colonel Black is this to Commander Red in Dragon Ball.
Dark Magical Girl Lutecia from Magical Girl Lyrical Nanoha StrikerS who cooperates with Mad Scientist Scaglietti not for his goals of world domination but to find and reanimate her mother, Megane, who is in a coma..
Pain in Naruto, and almost all of Akasuki in general. Also Sasuke.
Kabuto, because after Orochimaru was defeated by Sasuke, he ingrained some of his master's remains into his own body, and decided to help Madara in the 4th Ninja War.
Carrossea Doon from Madlax is only with Enfant to find and protect female lead Margaret Burton, who's been targeted by the group, rather than for Monday's insane plans.
Wei to Amber in Darker than Black. Wei serves Amber so that he can have a chance to fight Hei again
In the anime adaptation of Kirby, Meta Knight.
Nico Robin in One Piece starts out as a Dragon, but is later on revealed to have her own plans. After Crocodile's defeat, she joins Luffy's crew.
Saten in NEEDLESS, who is introduced as one of Simeon's Elite Four, yet in a few scenes he "accidentally" helped out the good guys as they infiltrated Simeon's HQ. It turns out that Saten was another survivor of the Adam Project who wanted Arclight dead so he could take over, but neither he nor his bosses counted on a revived Arclight coming back and absorbing him.
That's in the anime anyways.
Seto Kaiba from Yu-Gi-Oh! did this to his own Kaiba Corporation when he was what, twelve?
This trope is interestingly invoked in Neon Genesis Evangelion by Gendo Ikari, playing The Dragon to Keel Lorenz by ensuring the destruction of the Angels to open the path to Insturmentality. Of course, once the path is open, he will immediately take things off the rails for his own reunification scenario.
Dark Zero from Kurohime. Rather than follow the Big Bad Yashahime's plan and rule the universe, he just wants to exterminate humanity.
From Pokémon Special, Sird to Giovanni. Turns out she was The Dragon from another villainous team; she was actually working undercover as The Mole to learn more about various Pokemon from space.
In the original Mobile Suit Gundam, Char Aznable split his time between earnestly supporting Zeon's cause and trying to murder his boss's entire family in revenge for his father.
Char: Blame this on the misfortune of your birth.
Ichiki from Nabari no Ou plays The Dragon to Hattori before she betrays him for Fuuma. It appears that she'd been planning to dispose of Hattori all along, and was just waiting for him to outlive his usefulness.
Subverted in Gundam Seed. It's clear from fairly early on that Patrick Zala's right-hand man, Rau Le Creuset has his own agenda, though just what it is isn't made clear. Than The Reveal comes along: he's an Omnicidal Maniac who seeks the annihilation of the entire human race, and oh, by the by, is the real Big Bad of the series, with Patrick as his Unwitting Pawn.
In Saint Beast, Lucifer is initially this clearly having his own motives other than just serving Zeus, but eventually he rebels and tries to take everyone he can with him leading to a The Good, the Bad, and The Evil situation.
Gundam ZZ's Glemmy Toto starts out as a Villain of the Week, eventually graduating to a position as Haman Khan's right-hand man. While the two of them share the goal of recreating the glory of Zeon, Haman intends to do it herself. Glemmy, on the other hand, seeks to reinstate the Zabi dictatorship, as represented by himself.
Comic Books[edit | hide]
In the Green Lantern: Sinestro Corps War story arc, every major baddie had a different agenda.
The Anti-Monitor's goal is simple - to wipe out every single thing in the matter multiverse. Nice when a villain has such a clear focus on what they want, isn't it?
Sinestro was aiming to bring order to the multiverse by taking it over and ruling as the head of the Sinestro Corps in much the same way as he did his home planet of Korugar...or so he lets everyone think. In reality, his goal was not to win the war but to do so much damage that the Guardians of the Universe would allow the Green Lantern Corps to kill. In doing so, he believes he set in motion the events that would lead to the Green Lanterns being feared.
Hank Henshaw, the Cyborg Superman, is much cleaner, as all he has ever wanted was to die and be reunited with his dead wife. If the Anti-Monitor is Sinestro's Dragon, then Henshaw is the Anti-Monitor's as the Anti-Monitor promised him death. It almost worked. He was destroyed during the story but, in The Stinger, was located by the Manhunters and resurrected.
Superboy-Prime, meanwhile, only ever wanted to kill the Anti-Monitor as retribution for Earth-Prime being destroyed. He actually succeeded.
In Infinite Crisis, the main members of the Secret Society of Super-Villains:
Lex Luthor was apparently there to brainwash the heroes, but he had a secret pan to restart The Multiverse. And he would have gotten away with it too, if not for those meddling kids.
Talia al Ghul was there so that when the villains took over the world, she could betray he allies, kill the surviving villains and create an ecologically sustainable utopia.
Black Adam signed up to get villain help protecting his country. Even Bad Men Love Their Fatherlands.
Calculator and Deathstroke were in it for the money, fellows.
Doctor Psycho was... Uh... Ummm...
Mystique practically defines this trope. Her betraying the boss in charge is almost expected at this point, such as when she killed Mr. Sinister in a Batman Gambit to save Rouge's life.
Fanfic[edit | hide]
Past Sins has an interesting twist on the concept; the Big Bad is an extremely Reluctant Monster forced into the role, where The Dragon expects an Omnicidal Maniac.
The Doctor Who/Jem crossover fancomic Outrage reveals that Eric Raymond's toady Techrat was actually one of these.
In With Strings Attached, the Raleka have a completely different agenda than Brox and Co., but they have to work together to take the Vasyn pieces from the four.
In Ponies Make War, Nihilus serves King Titan with the intention of betraying him the moment she secures the Elements of Harmony. Princess Celestia anticipated this from the start, and actually helped to encourage it.
Films -- Animated[edit | hide]
Shenzi from The Lion King has an agenda. As the Matriarch of the hyena clan, who the Big Bad Scar used to keep the Pride Lands under control, she seemed to just want freedom, power and food for her clan.
Films -- Live Action[edit | hide]
In Con Air, Cameron Poe asks black supremacist Diamond Dog why he's content to be second in command to white Big Bad Cyrus Grissom. Dog replies that while his long term agenda is vastly different from that of Cyrus, he needs to escape before he can get back to it - going along with Cyrus is "a means to an end".
In Die Hard, Hans' top henchman Karl's brother—another of Hans' henchmen—is killed by McClane. Karl wants revenge against McClane, losing interest in the main mission for the rest of the movie.
The Twilight: Eclipse movie seems to do this to Jane, implying that she will allow the Cullens' deaths without Aro's knowledge. This makes more sense than in the book, where Aro wants Edward and Alice to join his side but nevertheless is willing to let Victoria's army kill them.
Literature[edit | hide]
Galadan Wolflord in The Fionavar Tapestry. Rakoth, the Big Bad, is a pretty standard dark god/ Evil Overlord who is out for world domination, but Galadan wants to destroy the world because he was driven mad by the death of his love, and he views Rakoth as a convenient weapon to help accomplish this. Obviously these goals are mutually exclusive, but it's implied that Rakoth knows all about Galadan's plans and does nothing simply because he knows the Wolflord isn't powerful enough to be a threat to him.
Per Word of God, Sauron only worked with Morgoth in The Silmarillion to advance his own ends- namely, to create perfect order by turning the world into a giant totalitarian state. As Morgoth's goals were, essentially, to take over the world so he could say "mine" before blowing it up, it's very likely that an Enemy Civil War would have broken out had the bad guys won. Fortunately for Sauron, Morgoth went down first, allowing him to graduate to Big Bad in The Lord of the Rings.
Grand Admiral Thrawn and Jedi Master Joruus C'baoth. C'baoth took Thrawn's orders with more and more reluctance as time went on, and he wanted very different things. Towards the end they weren't really on the same side anymore. Thrawn was fully aware of this, and at the time of his death was planning to solve the problem by eliminating C'baoth and making a replacement C'baoth. His own death, nearly simultaneous to C'baoth's, derailed his plan.
In the Codex Alera, Lady Invidia Aquitaine serves as her husband Lord Aquitaine's Dragon so that she can get her hans on the Realm's reins of power (and married him primarily for this purpose), while he wants to topple a First Lord he sees as corrupt, but has no real desire to hold power himself. After breaking with him, Lady Aquitaine winds up in a similar relationship with the Vord Queen, serving her only because she is able to keep Invidia alive despite her crippling injuries.
In Tad Williams' Otherland, John Dread is this to Corrupt Corporate Executive Felix Jongleur. Jongleur's Evil Plan is focused on living forever, and he's trained the Ax Crazy Serial Killer Dread as a Psycho for Hire to go after his physical opponents. Dread is far, far more ambitions than Jongleur realizes, however, and is simply biding his time, waiting for a moment of weakness to take over. And he does indeed, deposing Jongleur as master of Otherland and embarking on a godlike orgy of virtual destruction.
On the good guy side we get Vimes playing this role to Vetinari, maybe. Most of the time Vimes appears to be just a good little dragon, but he does sometimes have a different agenda, or at least a different motivation. Depending on how you look at it; technically, they both want what's best for Ankh-Morpork, but they do often differ on what exactly that means.
In David Eddings' The Malloreon, the Demon Lords Mordja and Nahaz work for Zandramas and Urvon solely so that they can seize the Sardion from them in their moment of triumph and then drag them into hell. Urvon's other Dragon, Harakan/Mengha is also a Dragon with an Agenda; he conspires with Nahaz to drive Urvon mad and take his place, so that the two of them can rule with the world. Since Nahaz is out to betray both of them, and both of them believe that their magic is enough to keep him under control, he succeeds in being Dragon with an Agenda and Dragon-in-Chief to two separate characters.
In Robert E. Howard's Conan the Barbarian story "The Phoenix on the Sword", Ascalante. He was summoned to help the conspirators, and in his first appearance, comments on how his tools think he is their tool.
Drake from Gone (novel). A psychopathic Complete Monster, he serves as Big Bad Caine's top enforcer, but it's apparent from his interior monologues that he's just helping Caine in hopes that Caine and Sam will kill each other. Although he probably would've turned on Caine as soon as they no longer had mutual goals, Caine tried to kill him after Drake gave Diana near-fatal injuries, which caused Drake to switch over to The Gaiaphage.
Live-Action TV[edit | hide]
In the 2009 V-remake S1 finale, we find out that Marcus definitely seems to have an agenda different from Anna's.
In Angel the incorporeal demon Sahjhan tried to use Holtz as his dragon by offering him vengeance against Darla and Angel. However Sahjhan's true goal was actually to either prevent the birth of Connor, Angel and Darla's unborn son, or kill him as a child. Sahjhan quickly grew frustrated with Holtz's focus on punishing the vampires instead of simply killing them as quickly as possible i.e. before Darla gave birth. And then Holtz decided that stealing Angel's son and raising him as his own would be a more fitting punishment, at which point Sahjhan totally lost control of his ally.
The Security Chief, who is Dragon to the War Chief's Big Bad in the Doctor Who serial The War Games.
In The Wire, it becomes apparent that Stringer Bell has very different goals from Avon, wanting to use the money from their drug operation to escape the ghetto and become a legitimate businessman. Avon meanwhile wants to protect his reputation and power as a gangster, and will even sacrifice smart business decisions to make war on the streets. This causes tremendous strain between them that builds until they both betray each other- Avon betraying Stinger to Brother Mouzone, and Stringer informing on Avon to the police. This leads to the downfall of the Barksdale organization.
Alex becomes this for Oversight during season 2 of Nikita
Tabletop Games[edit | hide]
The Ebon Dragon of Exalted serves as this to Malfeas. While he may advise Malfeas or defer to his authority, he constantly seeks to manipulate the Demon King to his own purposes, and endemically betrays him. One of the biggest exmples would have been back when they were the Dragon's Shadow and the Holy Tyrant respectively, and he manipulated him into creating the Unconquered Sun (a deity who could serve as something for the Shadow to oppose, even while he would also subvert the authority of the Tyrant).
In Warhammer, if you are a Dark Elf or a Skaven chances are you're this or have one or more of these under your comand. Wariors of Chaos and Orcs also might fit under this.
Similarly in Warhammer 40k, all Dark Eldar assumes this of their subordinates, as backstabbing is the only way to gain a promotion. Part of the reason why Incubi are valued as bodyguards over the Archon's own cabalites, Incubi loyalty is tied to their coffers, not your competency.
Video Games[edit | hide]
The man in the trope picture, Ocelot. In Metal Gear Solid, Metal Gear Solid 2: Sons of Liberty, and Metal Gear Solid 3: Snake Eater, Ocelot has been seemingly working for the Big Bad, but working behind their backs, and usually the backs of the people he's supposedly really working for. The portrayal of him in The Last Days of Foxhound even named the trope Chronic Backstabbing Disorder. The only MGS game he ISN'T this trope is the one where he's the Big Bad himself, Metal Gear Solid 4: Guns of the Patriots, where his agenda comes to fruition.
In Star Wars: The Force Unleashed, it is revealed that Darth Vader has been Palpatine's Dragon with an Agenda much earlier than the "I am Your Father" Scene in Episode V mentioned under Film. Vader tries pulling off a similar speech to the young Starkiller, but proves unable to sway him.
In the "Dark Side" DLC available for some versions of the game, Starkiller actually becomes a Dragon with an Agenda appealing to Luke Skywalker.
Astaroth in the Soul Series, according to the backstory.
In Suikoden III, you eventually learn Albert Silverberg orchestrated everything just to prove his skill as a strategist. He then casually derails it and walks away, likely leaving many players aching for a chance to settle the score. Or at least wipe that not-quite-smile off his face.
From the Paper Mario series, we have you guessed it; Dimentio. In this case, The Dragon is even more evil; an outright Omnicidal Maniac instead of a Woobie, Destroyer of Worlds.
The player was clued into the fact that Dimentio's trying to work behind Count Bleck's back pretty early on. However, it isn't until the end of the game that we learn that he's really higher up on the Sorting Algorithm of Evil, instead of just being a Reverse Mole.
Before that we had the Thousand Year Door where power hungry Sir Grodus tried to unearth an ancient power. Too bad said ancient power was a demoness whom Beldam served, and wanted to spring free. Grodus ends up living as a talking head afterwards.
Kefka from Final Fantasy VI kills the emperor at the end of the Disc One Final Dungeon and spends the next year killing whatever he pleases. He fits for this trope assuming he had an agenda in the first place.
All of Chaos' Warriors in Dissidia Final Fantasy have priorities other than serving him.
Cabadath, the Tall Man from the Chzo Mythos. The central conflict of the series, in fact, is him resisting replacement by Chzo. He fails in the last game, and the Player Character Theo becomes the new Dragon.
Dragon Age has Morrigan, who serves as The Dragon to Flemeth (though how evil Flemeth really is may be debatable), yet turns on her mother at the first sign of betrayal.
In Hordes of the Underdark, Mephistopheles spends the first two chapters manipulating events so that he can be freed of the Valsharess' control; culminating in the final battle of the second chapter in which he removes all obstacles between you and the Drow queen so that you can take his revenge for him.
The Ancient Minister and Ganondorf, as well as possibly Bowser, all take this role in Super Smash Bros. Brawl: Subspace Emissary
Hawke in the Advance Wars series. especially noticeable in Dual Strike as he actively chafes against Von Bolt taking control of Black Hole and eventually ends up leaving him for the heroes.
Samir Duran could be considered a Dragon With An Agenda compared to the Big Bad Infested Kerrigan in StarCraft when we discover some of his agenda in Dark Origin. He could also be viewed as an Enigmatic Minion and The Mole (against two different factions!)
Colonel Autumn from Fallout 3 counts, as both he and the President have different aims for Project Purity. He wants to use it as leverage to control all the free humans in the Capitol Wastelands, and President Eden wants to use it to exterminate everything in the Capitol Wastelands. It's also why he takes the opportunity to split as soon as the Lone Wanderer hits the Enclave base.
In Super Robot Wars Original Generation (as well as some non-OG games, to an extent), Shu Shirakawa acts as this to Bian Zoldark. Bian is well aware that Shu is both more powerful and more interested in pursuing his own goals, and he actually requests for Shu to merely observe during his final battle.
Cyril in Star Ocean 2
Kingdom Hearts: 358 Days Over 2 reveals that Saix and Axel were planning on overthrowing Xemnas together, and Axel had actually been following Saix's plan throughout Chain of Memories. Neither of them got as far as becoming The Starscream, however - Axel's friendship with Roxas and Xion led him away from the plan, and Saix apparently ended up dropping the idea and Becoming the Mask.
Xigbar also seems to have shades of this, looking back after Birth By Sleep, as he literally knows more about the backstory and mechanics of everything than the Big Bad, and is almost certainly manipulating his boss to an extent.
Vanitas displays hints of this in Kingdom Hearts: Birth By Sleep, by nearly killing a key member of his master's Xanatos Gambit before they've done their job, and allowing another person to live as a "plan b" in his own words suggests he has plans of his own, but in a subversion winds up following his master's plan exactly anyway.
Given how he is Made of Evil, it could be he didn't even have reasons for trying to kill Ven and letting the "plan b" live- he did it just because he could. And the novels reveal that his true motive is simply to rejoin with Ventus.
Mastema in Strange Journey. He outright states he hates the Three Wise Men and is just doing the job for the chance to ascend to a superior form.
Recurring villain Wild Dog in the Time Crisis series.
It is revealed midway through Assassin's Creed Brotherhood that Cesare, Rodrigo Borgia's Dragon (and son) is one of these, with plans to conquer all of Europe.
Which Rodrigo tries to fight, as while the Templars want world domination, they don't want to be seen doing it. They'd much rather work behind the scenes.
In Might and Magic VII, Kastore initially serves as an advisor to King Archibald of Deyja (along with the three other Terrans that followed his lead, but Kastore is the dominant one). Once you've made your choice between Light and Dark, however, it becomes increasingly clear that Kastore is building a power-base of his own (apparently going for a more traditional military-focused necromantic regime, but it turns out it is a bit more complex than that), culminating in Archibald evacuating his closest followers to a laboratory off the coast of AvLee and Kastore assuming the throne of Deyja.
Commander Shepard arguably falls into this role in Mass Effect 2, being forced to work with the Illusive Man stop the Collectors from abducting colonists. Ironically, the Illusive Man seems surprised when Shepard (possibly) opts to destroy the Collector Base, tells him where to shove it, before taking the Normandy and loyal crew for themselves.
The Black Knight in Fire Emblem: Radiant Dawn aided The Chessmaster in unleashing Ashera and her judgement and fought on their side in the game's climax, but didn't seem to have much interest in which side won in the end. His only personal goal was to surpass his teacher, Greil, as a swordsman. After Greil was killed he figured his son Ike would make a suitable substitute, and siding with Ashera gave him the chance to face Ike in combat. This goal also existed when he was serving Ashnard in Path of Radiance, but the player didn't know about it then.
Konishi from The World Ends With You. It's hinted at but eventually she outright betrays the Conductor by making a deal with Minamimoto, wanting to be the new Conductor when he becomes Composer.
Play a Horde character in World of Warcraft and go through the Mists of Pandaria scenario, and you quickly become this to Garrosh, eventually becoming The Starscream during the Seige of Orgrimmar.
Web Comics[edit | hide]
In Antihero for Hire, Angelson has at least one Xanatos Gambit going on under his boss's nose, we just don't know what.
Xykon and Redcloak have a mutually beneficial alliance in Order of the Stick. Redcloak only serves Xykon in order to open the Gates and get a fairer deal for the goblin race.
Although there's a fair bit more to it than that, as seen in Start of Darkness. Whether or not he'll turn Starscream eventually, or be forced to confront Xykon if their plans diverge, remains to be seen.
As it turns out, Redcloak's plan doesn't help Xykon's own goals at all, and Xykon is ultimately just Redcloak's Unwitting Pawn.
Also from OotS, Sabine is Nale's Dragon. However, she's also covertly working for The Inter-Fiend Cooperation Commission. Luckily for Nale, her agenda involves helping him advance his agenda.
And getting lots of hot hot sex from him.
Well, considering what she is, that last part goes without saying.
Gunnerkrigg Court: Ysengrin wants to Kill All Humans, and his boss Coyote apparently doesn't. It's possible that Ysengrin is trying to pull strings behind Coyote's back, and that Y killed Red and Blue because they saw too much.
Wanda from Erfworld seemingly serves Stanley the Tool willingly, but it turns out she convinced him to attack her kingdom of Faq because she expected him to get slaughtered, and when he emerged victorious, she only joined him in his quest for the legendary Arkentools because she was hoping she'd get to wield one herself. She does.
When asked why she's still with him after that, her answer is somewhere between Because Destiny Says So and You Can't Fight Fate.
Torg from Sluggy Freelance seems to be doing this professionally now. First with the Minion Master, and now with Crushestro ever since he found out that the Minion Master's sister betrayed him to Hereti-Corp.
Ian and Anita in Errant Story fit this trope pretty well, with the added little twist that it isn't totally clear which one is the Big Bad and which one is The Dragon.
Web Original[edit | hide]
In The Gamers Alliance, Ronove becomes the closest lieutenant of Duke Malphas after the siege of Vanna but has his own, secret agenda for working with the demonic duke.
In the That Guy With The Glasses world, The Other Guy is the real puppet-master behind the site. The one who everyone thinks is the ineffectual Bad Boss, The Nostalgia Critic, is just like that because he's desperate for attention and praise.
Linksano and Mecha-Kara are ultimately this during the Lord Vyce arc of Atop the Fourth Wall. In fact, Linksano defects to the side of the good guys to take Vyce down, while Mecha-Kara was a former Big Bad who only went along to take out Linkara.
Western Animation[edit | hide]
Destro in some iterations of G.I. Joe. He's pretty much in it for the money and The Baroness.
In Teen Titans, Slade (formerly a Big Bad himself) only serves as Trigon's Dragon so he can get his mortal body back. Interestingly, he starts out this plotline very subservient to Trigon, and gradually starts acting less as a servant and more like a partner when he gets closer to getting what he wants. When it becomes apparent that Trigon has no intention of honoring their agreement, Slade teams up with the Titans to take him down.
Raythar from WITCH serves this role during the first part of the second season- Big Bad Nerissa has her own agenda, but Raythar just wants to avenge the previous Big Bad, Phobos. When a chance arises to put Phobos back on the throne, he ditches Nerissa and transfers his loyalty back to his original boss. Nerissa, meanwhile, replaces him with Shagon, who is her creation and therefore completely loyal.
The Fright Knight in Danny Phantom subtly expresses his absolutely distaste working for Pariah Dark and it hints he wants to go a different route. It ultimately proves true when he works against Pariah during the final battle. It's hinted he went to serve Vlad, but the original writer left the show before possible stories of this could have been told in Season Three.
On Gargoyles, the Weird Sisters serve this role for the Archmage—he wants revenge on Goliath, while they want revenge on the Magus and the rest of the Avalon Clan. They couldn't really get on the island themselves, however, since Oberon had forbidden it; the reason they willingly put themselves under the Archmage's service was because they knew he would order them to go there, exploiting a loophole in Avalon's laws.
They could arguably be considered this for Oberon later, though that depends on whether or not he counts as a villain. Word of God says they are still plotting revenge, despite Oberon being at peace with the Avalon gargoyles.
Sonic the Hedgehog Snively in the final minutes of the series finale was revealed to be this.
In the original My Little Pony, Porcina's Raptorian minions encourage her to go to the extreme of turning Ponyland and all the Little Ponies into glass because they want to take the place for themselves.
The X-Men: Evolution version of Mystique. In the first season, she's working for Magneto, but they have a slight falling out and works independently for the rest of the series. This causes some tension when both tried to take command of the Brotherhood, with varied results depending on the episode and the Brother.
In a dark future episode of The Grim Adventures of Billy and Mandy, Grim is ultimately this to dark overlord Mandy. She was aware of this and set things up to capture the rebels and make Grim pay.
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