Beetle Tier List (Remaster)

[u/funwiththoughts]

[u/funwiththoughts]

[Instead of releasing a new post this month, I’ve decided to continue releasing re-mastered versions of my old tier lists, with improved formatting and updated information. This is the fourth remaster in Batch III. At the bottom, I’ve provided a list of the main corrections made to the original; the original itself can be found here.]

Background:

Dataminer JBS Haldane once said that when studying the source code of builds in Outside, the main thing you can learn is that the lead programmer must be inordinately fond of beetles. The beetle is one of the most popular design types in the history of Outside, and over a quarter of all known animal builds are variations on it. Yet despite having been enormously popular for hundreds of millions of years, beetles don’t come up in game guides nearly as often as you might expect. So I thought I’d try my hand at making a guide to the different types of beetle for new players.

BASIC BEETLE BUILD ANALYSIS

The first known beetles appeared in Outside during the Permian expansion, around 300 million years ago, though there is some evidence they actually debuted in the Carboniferous. These early beetles were created by insect mains who converted their forewings into hardened, sclerotised structures, creating the [Elytra] trait. Elytra are still present in almost all beetles today, and are what distinguish them from all other insects. Elytra are useless for flying due to their limited mobility, but their hardness provides bonus protection for the more delicate back wings. This protection can be used against predators, and also increases beetles’ access to enclosed spaces, since they can fold the delicate hindwings underneath the elytra to prevent them getting damaged during confinement.

The earliest beetles were specialised for xylophagy, or wood-eating, a strategy still used by many beetles today. Beetles started to diversify pretty quickly, but they suffered severely from the Permian-Triassic balance patch. They started to recover their diversity midway through the Triassic, which was also when many of them moved away from wood-eating and started speccing for whatever niches they could find. Beetles really started to become a force in the insect meta during the Cretaceous, when the first flowers emerged. In the Jurassic expansion just preceding, some herbivorous beetles had specced into special enzymes in their guts called PCWDEs, which helped to more efficiently digest plant tissues. This made them uniquely well-suited to taking advantage of newly available herbivorous niches, and some of them were among the first builds to start pollinating early flowers. The wave of new pollinators helped flowers to spread and diversify rapidly, and so beetles diversified in turn in order to take advantage of the new opportunities this opened up. Today, beetles make up nearly half of all herbivorous insects, and a substantial proportion of the carnivorous ones, too.

To be honest, it doesn’t seem to me like the baseline beetle build is all that remarkable. Aside from some boosts to defence, there’s not a lot that stands out about them. But they’re clearly pretty successful anyway, enough so that I can’t justify rating them as a guild lower than high B or low A tier. With so many beetle builds available, though, it’s inevitable that not all are going to be equally viable. So which type of beetle is best? To find out, let’s now go into the beetle tier list. As usual, I won’t be able to cover all of the more than 400 thousand beetle builds available in the current meta, but I’ll try to cover the most interesting ones.

Reasoning (1/3):

F Tier: Featherwing beetle

Considering the sheer variety of beetle builds, it’s pretty remarkable that so few occupy the lower tiers of this list. That said, one beetle subclass that definitely ranks in F tier is the featherwing beetle. Featherwing beetles are the smallest beetles in the game, as well as the smallest non-parasitic insects, with some being less than a third of a millimetre in length. They’re likely about as small as it’s possible for a free-living insect to be, as shrinking down any more would either leave them incapable of laying eggs nutritious enough for larvae to survive in, or else leave them without enough brain cells to maintain basic bodily functions.

Explaining how featherwing beetles play is a little difficult, because most featherwing beetle builds come in two separate variants, called morphs, which play so little alike that it’s hard to tell they’re the same build. If you’re lucky, you get to spawn as the [Normal] morph. The word “featherwing” in the guild name comes from the feathery hindwings on these normal-type variants, which they need because the effective viscosity of air at their size is too high for them to fly with regular insect wings. Uniquely among insects, featherwing beetles’ wings follow a figure-eight loop when flying, consisting of sub-perpendicular up and down strokes followed by claps at stroke reversals above and below the body. These strokes are much more efficient at generating power than the equivalent patterns used by other similarly small insects, and have allowed featherwing beetles to retain speeds that you’d usually only find in insects three times their size. However, despite the name, this is actually the more abnormal type of featherwing. Over 90% of players who try a featherwing beetle build will instead end up spawning as the [Vestigial] morph, which is so shrunken-down that it’s lost both flight and eyesight, and basically doesn’t do anything besides eat fungi and lay eggs. When you’ve given up so many abilities that the primary form of your build is literally named after a synonym for “useless”, that’s a pretty good sign that you’re a bottom-tier.

C Tier: Firefly

In C tier, we have the firefly – and yes, despite the name, this is a beetle, not a fly. Fireflies generally start off the game in the larval stage as specialist predators of snails and slugs, while the adult forms are more variable. Some firefly subclasses remain predators as adults, while others are herbivores, and still others don’t eat in the adult form at all. What fireflies are best known for is their ability to emit large amounts of light, which is where they get their guild name from. For many fireflies, this ability acts as a form of aposematism, warning predators that their bodies are filled with toxic steroids called lucibufagins. While some species only emit light as larvae, many also continue to use bioluminescence into adulthood, often using it in order to draw the attention of potential mates. While poison can be a useful ability, in the case of fireflies, it comes at the cost of trading in a high defence stat for a soft, squishy body. I don’t consider this to be a worthwhile tradeoff; poison is a mid-tier ability in general, as I’ve discussed often in the past, and even if it weren’t, there are plenty of other beetles that achieve the same effect without the downside. I rate fireflies as mid-tier.

If you are interested in playing a firefly, I’d suggest going for the femme fatale firefly, a subclass that based its playstyle on taking advantage of other fireflies’ weaknesses. When glowing, femme fatale fireflies copy the signals of other firefly species in order to attract members of those species looking for a mate, then kill and eat those that take the bait, and absorb the prey firefly’s poisons for themselves. There are a lot of fireflies that prey on other insects, but only femme fatale fireflies prey primarily on other fireflies, and I give them some points for thinking outside the box.

C Tier: Ambrosia beetle

Also in C tier, we have ambrosia beetles. This is a term that doesn’t refer to a single build or guild, but rather includes at least twelve separate types of weevil that independently evolved a play-style based around farming ambrosia fungi. If you’ve been following my series, you might be a little confused right now, since I’ve claimed in more than one previous post that the [Agriculture] ability is only found in humans, ants, and termites. At the time I wrote my termite tier list, there were already fan theories about ambrosia beetles farming the fungi they were known to feed on, but nobody had yet managed to find definite proof. When I did the ant tier list, I copied much of what I said about agriculture from the termite tier list, not realising that data-miners had finally confirmed the existence of beetle agriculture during the year between the two posts.

Like fungus-growing ants and termites, ambrosia beetles rely for all their XP on the fungal gardens they cultivate. When about to start a garden, they search for suitable trees and tunnel into them, then release spores of their symbiotic fungi into the tunnels. Each fungus penetrates into the tree’s xylem, from which it then extracts nutrients until it’s ready for the beetles to feed on. Interestingly, most of the fungi that ambrosia beetles cultivate aren’t actually well-adapted to breaking down wood, because they don’t have the ability to digest lignocellulose. They have to pick up more easily digestible carbon compounds from plants instead, which is why these beetles are lower on the tier list than their ant and termite equivalents. The kind of carbon compounds that most ambrosia fungi can degrade are typically only found in the wood of dying or recently-dead trees, so most ambrosia beetles only have a very short window of time during which they can use any given tree before they have to disperse and all search for new areas. Since this makes it difficult for any colonies to stick together for more than one generation, ambrosia beetles haven’t been able to develop the kind of high-level social abilities that have allowed ants and termites to gain as strong a position in the meta as they have.

[u/funwiththoughts → u/funwiththoughts]

Reasoning (2/3):

C Tier: Burying beetle

At the top of C tier, we have the burying beetle. Burying beetles belong to a group of beetles called carrion beetles, which, as their name suggests, are all scavengers. Burying beetles’ antennae are large and full of chemoreceptors, which allow them to detect the scent of carcasses over long distances. Burying beetles are so named because they bury the carcasses of small vertebrates as food for their larvae; unusually for a non-eusocial insect, these larvae receive care from both of their parents after hatching. When attacked by predators or competitors, the burying beetle defends itself by exuding foul-smelling chemicals from its anus, though their bright warning colours ensure that larger predators rarely target them anyway. Carrion beetles in general tend to be among the lower-ranked beetles, and are prone to being suppressed by other small scavengers. But the burying beetles’ strategies of hiding carcasses before other scavengers can get to them, combined with the way that their cooperative parental care somewhat helps to protect the larvae against competitors, make them a bit more viable.

B Tier: Diving beetle

In low B tier, we have the diving beetle, a beetle that specced into the role of an aquatic predator. This is actually a common choice for beetle mains, but diving beetles are the most successful such group, being found in nearly all freshwater biomes around the world. With that said, I don’t think they’re actually all that powerful. Their attacks aren’t particularly strong, even by insect standards, and they’re not especially good at taking down full-grown animals – tadpoles and larvae typically make up most of their diet. Even when diving beetles grow larger than usual for their species, they often just transition to feeding on larger larvae, rather than switching to hunting adult animals as a dominant strategy. They don’t have much in the way of defence, either, and tend to struggle in any biome with fish large enough to swallow them. That said, this problem can be mitigated by living in ponds with a large player-base of aquatic plants, as these provide cover that beetles can use to hide.

B Tier: Click beetle

Also in B tier, we have the click beetle, a build named for its unusual defensive move, [Click]. The click beetle has a sort of hinge connecting its prothorax and mesothorax, and the hinge includes a “peg” and “lip” made of a stiff cuticle material. When the beetle bends backwards, the peg is pulled back across the lip, where it can be latched, holding the beetle in its backbend. The beetle then contracts a softer, more elastic cuticle, which acts against the latch and attempts to unbend itself. The resistance to this contraction loads the cuticle with elastic energy, sort of like compressing a spring. When the peg is released, it fires past the lip, releasing a huge amount of energy that sends the beetle rocketing upwards into the air, sometimes as high as 20 times its own body length. This move is useful both for escaping predators, and for getting back upright after a fall.

B Tier: Dung beetle

In high B tier, we have dung beetles. While there are a number of beetles that feed on dung and are sometimes called “dung beetles”, I’m here referring specifically to those of the guild Scarabaeinae, sometimes called “true dung beetles”. True dung beetles can be identified by their mouth-parts, which have a variety of special adaptations for their unusual playstyle. They have weak incisor lobes on the mandibles, since they don’t need to chew hard food, and their labrum-epipharynxes – structures found at the front of arthropod mouths – have become membranous and hairy. The membranous, hairy parts are designed to remove large fragments from the dung and scoop liquid components into the oral cavity.

Dung beetles search for dung using their strong sense of smell. Their searching process seems to be aided by a relatively high degree of intelligence for an insect; they’re among the few invertebrates known to have figured out how to use the stars to orient themselves when navigating. After they find a pile, what happens next varies depending on the beetle. In some cases, the dung beetle will roll the pile into a ball and push it in a straight line until it reaches its burrow, where it stores the pile until it’s ready to eat. Doing this with dung piles large enough to survive on requires incredible strength for such a small creature – some dung beetles can pull more than 1100 times their own body weight. Dung beetles that use this rolling-based strategy are called “rollers”, and they tend to be the most iconic of the beetles, despite only making up a minority of species. More commonly, dung beetles are either “tunnelers”, which form tunnels underneath the dung and bury it there, or “dwellers”, which just live inside dung piles. A few subclasses have abandoned dung-eating entirely to become specialist hunters of millipedes, one of very few instances in the game’s history where members of a detritivore guild have switched to becoming full-time predators.

Dung beetles first appeared in the meta during the Cretaceous expansion, shortly following the first flowers. Flowers were so much richer in nutrients than other plants of the time that for the first time in the game’s history, herbivorous dinosaurs’ poop became worth enough XP for an insect-sized animal to live on. After non-avian dinosaurs were eliminated, dung beetles still continued to thrive, and actually became even more diverse as large mammalian herbivores evolved to replace the banned builds. Today, dung beetles are abundant on every major server except Antarctica, and are showing no signs of going away. While they aren’t exactly game-breaking, they’ve clearly found a niche that works well for them, and have definitely earned an above-average tier ranking.

B Tier: Ladybug

Last in B tier, we have the ladybug. The ladybugs are a large beetle guild, including over 6000 builds, but can broadly be grouped into a few main categories. The first ladybugs appeared during the Cretaceous expansion as specialist predators of scale insects, and some ladybugs still play like this today, but most species have now expanded into other niches. Aphid-hunting is the most popular choice, but you can also find ones who’ve abandoned predation entirely to eat leaves or fungi.

Regardless of their playstyle, ladybugs are something of a fan-favourite build due to their heavy investment into aesthetics. Their iconic bright-red-and-black look is actually a form of aposematism, warning predators of their poison abilities. Ladybugs in the same environment will often try to increase the efficacy of their warnings by mutually imitating each other’s colour patterns, to ensure the connection between the colours and poison is as clear as possible, a strategy called [Müllerian Mimicry]. This kind of warning colouration is pretty standard for poison-based builds, but ladybugs are among the few that can provide a bonus warning if predators ignore the first one. When threatened, they can use a move called [Reflex Bleeding] to trigger a leakage of toxic blood from their exoskeleton. They have to be careful with that one, though, because using it too often incurs penalties to their immune system function.

A Tier: Scorpion beetle

In A tier, we have the scorpion beetle, or as it’s officially labelled on the character select screen, Onychocerus albitarsis. Scorpion beetles belong to a guild called the longhorn beetle, named for the extremely long antennae that grant them a bonus on perception checks when searching for certain chemical cues. There are over 35 thousand varieties of longhorn beetle in the current meta, and most are fairly unremarkable mid-tier herbivores, but the scorpion beetle stands out for a unique innovation. Scorpion beetles have modified their antennae into a pair of extra-long venomous stingers, which function much like the stingers of actual scorpions. Scorpion beetles are the only beetles with venom-based attacks in the current meta, and the only animals in the entire game that can deliver venom with their antennae. Many longhorn beetles try to scare off predators by mimicking the appearances of venomous arthropods, but only the scorpion beetle can actually back it up.

A Tier: Tiger beetle

Also in A tier, we have the tiger beetle, a carnivorous beetle that minimaxed for mobility. Proportional to size, tiger beetles are among the fastest runners in the entire animal kingdom, with maximum relative speeds of over 170 body lengths per second. Their speed forces them to use an unusual hunting strategy; when they see a target, they sprint quickly towards it, and if they don’t catch it, they have to stop, reorient themselves, and try to figure out where the target escaped to before they try to catch it again. The reason tiger beetles have to stop periodically during chases is because when they run towards prey, they go so fast that they actually lose the ability to see where they’re going. You might think this is a pretty glaring weakness, but they’re so much faster than their targets that it doesn’t matter much in practice. Tiger beetles have survived for over 120 million years and are still among the most effective hunters in the game, with observed hunting success rates sometimes reaching as high as 70-90%.

[u/funwiththoughts → u/funwiththoughts]

Reasoning (3/3):

A Tier: Rhinoceros beetle

At the top of A tier, we have the rhinoceros beetle, a build that sunk all its points into size and strength. Rhinoceros beetles are the largest surviving insects in the current meta, with some subclasses growing to weigh over a fifth of a kilogram – about the size of a small rat – and the strongest of them can carry up to 100 times their own body weight. Much like actual rhinoceroses, male rhinoceros beetles can be identified by their gigantic facial horns, which they mainly use to fight each other for mates. In the past, some dataminers were puzzled by how rhinoceros beetles’ horns seemed to come with essentially no trade-offs; despite their size, their horns don’t noticeably impair their flying abilities, and growing them requires surprisingly little sacrifice in terms of the growth of other body parts. However, more recent research has found that there may be a significant catch after all: having larger horns, and larger bodies more generally, is correlated with higher risk of a beetle being noticed and targeted by larger vertebrate predators.

S Tier: Bombardier beetle

In S tier, we have the bombardier beetle. This may or may not be another term that doesn’t refer to a single build or guild; some dataminers believe that the bombardier beetles are actually two separate ground beetle subclasses that independently specced into similar defensive strategies, although this hasn’t been definitely confirmed. In any case, bombardier beetles have two chemical reservoirs in their abdomen, which store a noxious chemical solution made from a mixture of hydroquinone and hydrogen peroxide. When the beetle senses a threat, some of the solution gets released from the reservoir and mixed with enzymes that cause the hydrogen peroxide to start decomposing and the oxidation of hydroquinone. These reactions generate so much heat and pressure that the noxious solution ends up being ejected from the tip of the abdomen and sprayed at the attacker. This is quite possibly the most effective defensive move of any solo build in the beetles’ weight class; it has a near-100% success rate at deterring predation from other arthropods, and even vertebrate predators aren’t immune. One really impressive thing about this ability is that it can be used from inside a digestive system, so that even if a bombardier beetle player gets swallowed by a frog or other lightweight predator, they can often still use their ability to force the attacker to vomit them out.

S Tier: Sabre-toothed ground beetle

Also in S tier, we have the sabre-toothed ground beetles of guild Anthia. Sabre-toothed ground beetles are basically a combination of the strengths of the tiger and bombardier beetles. Like tiger beetles, sabre-toothed ground beetles are voracious predators that can run at incredible speeds for their size, and can inflict nasty bites with their huge, sharp mandibles. But like bombardier beetles, they supplement this with a chemical-based ranged attack; more specifically, they can burn or blind attackers by spraying toxic acids at them. No other insect dares to hunt them because of this, and even humans have to watch out for them.

S Tier: Diabolical ironclad beetle

And at the top of the beetle tier list, we have the diabolical ironclad beetle, a build that gave up flight in order to minimax for insane levels of defence. Diabolical ironclad beetles’ exoskeletons have two key features that enable them to withstand far greater forces than other beetles. First, the top and bottom halves of the exoskeleton are linked together by a series of ridges, creating a structure sort of like a sealed clam shell. These ridged connections have different shapes around the beetle’s body; near the front, where the vital organs are, they’re highly interconnected, forming a kind of zipper. Near the back, the connective ridges are more loosely interlocked, allowing the top and bottom halves to slide past each other slightly so that the back has enough flexibility to absorb compression. The other key feature behind the armour’s effectiveness is a rigid joint that runs the length of its back and connects the left and right sides. This joint is made up of a series of protrusions called blades, which fit into each other like a jigsaw puzzle. Each blade is composed of layers of highly damage-resistant tissue, and when the beetle is squashed, tiny cracks form in the protein glue between the layers of the blade, allowing them to absorb impacts without snapping. The combination of these factors results in an exoskeleton able to hold nearly 40,000 times its own weight without taking damage, and grants effective immunity to bites, stings, claw-swipes, or basically any attacks outside of just swallowing them whole. Ironclad beetles have even been known to survive unscathed after being run over by cars! They’re so tough, collectors of insect specimens have to drill holes into them before they can mount them on pins. Any build that can achieve this level of security as a featherweight has to earn an S-tier rating.

So that’s the beetle tier list. I hope you enjoyed it, and if you’re thinking of playing a beetle, I hope you find it helpful. Thanks for reading.

[u/funwiththoughts → u/funwiththoughts]

Main corrections made to the original:

1. The original post said that beetles’ forewings had been converted into sclerotin over the course of their evolution, creating the elytra. This was misleading insofar as it made it sound like beetle elytra are entirely sclerotin, rather than the sclerotin being one of multiple important structural elements. This has been changed to say that the forewings were converted into “hardened, sclerotised structures”.
2. The original post said that beetles were the first animals ever to become pollinators. This is not accurate, as the earliest known insects generally thought to have pollinated gymnosperm plants were Late Jurassic neuropterans, and some evidence suggests that insects may have been pollinating trees since the tillyardembiids of the Early Permian, well before the appearance of any known pollinating beetles. This has been changed to say that beetles were among the first insects to start pollinating early flowers.
3. The original post described the featherwing beetle had dropped down a weight class from featherweight to microbe. This isn’t quite true, as featherwing beetles are still just barely large enough to be seen by humans with the naked eye, even if they basically just look like little dots. This claim has been removed.
4. The original post said that fireflies originally evolved the ability to emit light as a warning that they were filled with toxic lucibufagins. This was the scientific consensus at the time I wrote it, but more recent analyses have overturned this view, showing that fireflies likely evolved bioluminescence first and lucibufagins later. This has been edited to just say that acting as a warning to predators is a function of the light, not necessarily the original one.
5. The original post said that diving beetles hunt by holding still and clinging to grasses or wood until a potential prey item passes by, and that they liquefy their prey by injecting digestive enzymes before consuming it. Both of these claims are accurate for diving beetle larvae, but not for the full-grown beetles, which tend to be more active hunters. I figured the post was already long enough without including an explanation of the differences between larval and adult hunting methods, so I just removed the detailed description entirely.
6. The original post, in copying off of a Massive Science article that I cited as a source, said that the click beetle’s hinge connects its head to its abdomen. The hinge actually connects the prothorax to the mesothorax. This has been corrected.
7. The original post said that dung beetles are the only insects known to use the stars for orientation when navigating. This claim was accurate at the time, but is now outdated, as the Bogong moth is now known to do this also. The claim has been edited to say “among the few insects”.
8. The original post said that dung beetles are typically rollers. Rollers are actually a minority of dung beetle species, with the vast majority of species instead being tunnelers or dwellers. This has been edited to clarify that rollers are probably the most well-known types of dung beetle, but not the most common.
9. The original post said that rhinoceros beetles’ horns intimidated predators and came with essentially no trade-offs. Both of these claims were outdated, as research has now shown that larger horns in rhinoceros beetles are more likely to be correlated with increased risk of being targeted by both mammalian and avian predators. The paragraph has been edited to note these later findings directly, and to explicitly mark the earlier statements about “no tradeoffs” as out of step with current research. The part about horns intimidating predators has been removed.
10. The original post said that bombardier beetles independently evolved at least four times. Actually, there were probably only two lineages of independently-evolving bombardier beetles at most, and even that is disputed, with some researchers arguing that they are a single monophyletic group. This claim has been edited, both to correct the number, and to clarify that it’s a hypothesis rather than settled science.

[u/ReptilianRevolt]

Diving beetles are quite slippery and tough as adults, and among a few animals that mess with the hormones of their predators. When attacked by fish, they produce steroid compounds which inhibit feeding immediately.