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I want a world in which animals roam the wilderness on four legs, yet at least some of them are able to do the kind of carrying and fine handling of objects done by humans. The planet is superficially similar to Earth, but it is not Earth, and there are no humans around (at least not yet; who knows what humans might do once they figure out interstellar spaceflight...)

There seem to be basically two ways to go about reaching that goal; either give the ancestor of land life on that planet six legs, allowing for two to evolve into arms and hands in a manner similar to how those of humans evolved; or give the ancestor four legs, and have their forelimbs serve both purposes, not unlike gorillas on Earth.

For the alien feel, as well as the additional options it gives, I'm currently leaning toward six legs, the foremost two of which could evolve (or not, depending on the species) into hand-like extremities.

This is intended to be a realistic world, so the normal issues of evolutionary selection pressure apply.

Suppose that an intelligent designer is faced with the choice of, for an ancestor species, ticking the "four legs" or the "six legs" checkbox on the requisition form. What, if any, would be good reasons for them to select four legs rather than six? Or, in other words, what disadvantages would be confered to the creature by having six legs as opposed to four?

For simplicity's sake, you may assume a single ancestor species for all relevant land life on the planet, so for example "some other species would be more energy-efficient in not needing to grow an extra two limbs" does not apply. Other species on the planet may have a different number of legs (compare to how there's also spiders and caterpillars on Earth), but can be ignored for the purposes of this question.

This question is basically the opposite of Why would an animal need six legs?

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    $\begingroup$ On Earth we have both (not to mention 8 and ten legged critters, from distinctly different lineages of animals. All arthropods have 6 or more legs, exoskeletons, open circulatory systems, and no spinal cord. All chordates have 4 or fewer limbs. There are significantly more arthropods by weight than chordates in the world today. $\endgroup$ – pojo-guy Sep 27 '18 at 17:51
  • $\begingroup$ @pojo-guy You're right about Earth having both (and more), but often, those don't compete in the same weight class. I'm thinking more in terms of the same species or at least group of species, with the main difference being the number of legs: would four legs give a clear advantage over six? $\endgroup$ – a CVn Sep 27 '18 at 17:54
  • $\begingroup$ You only need 3 legs to stand firmly and effortlessly, and symmetry for easy walking, that gives 4 (species with 2 can't sleep standing, because it requires active balance control, and animals who lest one leg usually can still run, but it seems more difficult to them). So everything above 4 seems a cost to grow and power up, with little to no functional benefit. Seems - I can't find sources decent enough to write an answer now. $\endgroup$ – Mołot Sep 27 '18 at 18:09
  • $\begingroup$ So, if I understand correctly, you're asking for six limbs but a quadrupedal gait, like a centaur? $\endgroup$ – SealBoi Sep 27 '18 at 18:15
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    $\begingroup$ @Mołot: “A bird’s wing, comrades,” he said, “is an organ of propulsion and not of manipulation. It should therefore be regarded as a leg. The distinguishing mark of man is the HAND, the instrument with which he does all his mischief.” The birds did not understand Snowball’s long words, but they accepted his explanation, and all the humbler animals set to work to learn the new maxim by heart. FOUR LEGS GOOD, TWO LEGS BAD,... Animal Farm $\endgroup$ – nzaman Sep 29 '18 at 6:15

13 Answers 13

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There's at least one paper suggesting the reason for four limbs is because of the way embryos develop.

Basically, an embryo's cells develop early on into three layers: ectoderm, mesoderm, and endoderm. The endoderm becomes the digestive tract, the ectoderm the skin and nervous tissue, and the mesoderm everything else (bone, muscle, other organs). The mesoderm itself splits into two layers, one that lines the inside of the body cavity, the other that lines the gut.

The hypothesis is that the places the mesoderm will form bone and muscle that creates limbs is near the front and rear ends of the gut, because that's where the layers are close enough together to interact with the ectoderm and each other, but not close enough to merge completely, which leads to the formation of the tail and head, and not far enough apart, as they are along the gut, to prevent the tissue that will become limbs from forming.

If that's true, then in order to have six limbs, you need the early gut, for some reason, to be shaped kind of like an hourglass or peanut, some place where you get the right conditions for the proper distances to allow for limb creation between the front and rear ends. If this hypothesis is true, the the choice between four limbs and multiple limbs is set very early on in development, long before a body would have to deal with allocating resources for limb development.

That being the case, there's no particular reason why six limbs wouldn't develop if the early embryonic development wasn't just a slight bit different from ours.

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    $\begingroup$ that doesn't work, insects have six limbs and use the same hox genes as us. $\endgroup$ – John Sep 27 '18 at 22:17
  • $\begingroup$ I merely report what the paper suggests. $\endgroup$ – Keith Morrison Sep 27 '18 at 22:51
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    $\begingroup$ @John Keep in mind that insects are invertebrates and employ exoskeletons. $\endgroup$ – A Lambent Eye Jan 15 at 13:33
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    $\begingroup$ They use the same 3 developmental tissues. The paper is full of errors, the front limbs don't even form at the end of the gut, its just bad science. $\endgroup$ – John Jan 15 at 13:43
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The only disadvantage that I can see is the extra consumption of resources.

We can see from Earth that six legs are not needed. Therefore the extra resources that are needed to grow and operate those extra legs could be put into other things (other adaptations or a slight increase in overall population).

I don't see this as a huge advantage but it is the only thing that I could think of that would answer this question.

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    $\begingroup$ The larger size of shoe shops alone would cause the economy to skew drastically. $\endgroup$ – VBartilucci Jan 14 at 18:38
  • $\begingroup$ Of course many many organisms get stuck with suboptimal body plans becasue they inherit them. Evolution can only work with what you have after all. $\endgroup$ – John Jan 14 at 22:21
  • $\begingroup$ I think it necessary to point out that "extra consumption of resources" is a serious matter. In nature, scarcity is the norm: most animals will die to either starvation or predators, so anything that costs additional energy to maintain will be weeded out ruthlessly by evolution unless it provides a clear advantage to compensate. Two more limbs is a huge cost just in materials and maintenance (never mind any other factors), which requires a larger food supply: the extra two limbs must have a hugely significant purpose if they are to outweigh that massive cost and so not be selected against. $\endgroup$ – Palarran Jan 15 at 2:18
  • $\begingroup$ @John, exactly, evolution chooses between two options. If the other choice doesn't show up or only shows up with other disadvantages, the change won't happen. $\endgroup$ – ShadoCat Jan 15 at 19:32
  • $\begingroup$ @Palarran, the extra consumption of resources is serious, as you say. However, change only happens if random mutation produce a difference that is more advantageous than disadvantageous. There is also a matter of luck: Super Mutant Rodent got stepped on by a triceratops before it could breed. $\endgroup$ – ShadoCat Jan 15 at 19:43
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There is a very real disadvantage to more appendages.

Controlling more appendages simultaneously takes much more concentration. Ask any fingerpicking guitarist. If you watch them they don't use the little finger on the picking hand. Because it's extremely difficult to control when you're already concentrating on the rest. Going from using 3 to four is relatively easy. 4 to five isn't.

So coordination of extra limbs would be a big factor in fine control scenarios.

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    $\begingroup$ Interestingly enough, the exact opposite argument also works well. One advantage of 6 legs for insects is that they require much less control because it's easier to create a stable base. Its only once you have the brain power to stand up on fewer legs that you start wanting to control them more precicely. $\endgroup$ – Cort Ammon Sep 27 '18 at 23:44
  • $\begingroup$ @CortAmmon I'm not sure how insects manage it, but would hardly call them sentient.... complex tasks requiring concentration are different from innate ones that don't need much, there is a whole lot more to fingerpicking guitar than just controlling your hands, and you're not going to be doing it while running. $\endgroup$ – Kilisi Sep 28 '18 at 0:13
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    $\begingroup$ There's two solutions to the problem. one is to develop the musculature and control required to fingerpick. The other is to evolve in such a way that you don't need to fingerpick. Insects took the latter path. $\endgroup$ – Cort Ammon Sep 28 '18 at 0:17
  • $\begingroup$ @CortAmmon no idea what you're on about mate, feel free to downvote and compose your own answer $\endgroup$ – Kilisi Sep 28 '18 at 3:25
  • $\begingroup$ That's not a real disadvantage, brains evolve to have enough capacity to distribute to all the limbs, picking is using your limbs on a way they did not evolve for. that's like saying having fingers is a disadvantage to humans because typing requires concentration. $\endgroup$ – John Sep 29 '18 at 5:06
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Interesting question, Sleipnir approves. The short answer is, it sort of depends on your creature's body design.

From the reading it sounds like you are describing something that would function similarly to a centaur...possibly. It's also possible that the creature would travel on all six legs with the front pair functioning similar to a gorilla where they can be used for both travel and more fine motor skills.

The finer you want the motor skills to be the less likely your creature will use the front pair of limbs for locomotion.

What your creature is specialized for could range pretty widely as well.

I can envision a prairie/grassland creature with bison or rhino like characteristics using its size and bulk as a defense...maybe some horns just for fun.

  • This creature would be low to the ground with strong legs. It would be very stable.
  • The complications of extra legs in the creature's gait could be removed/mitigated by short stout legs and a slightly longer body than traditional prairie creatures
  • This body design doesn't work particularly great for giving it precision digits, if you want to do that you probably need something more akin to a centaur body design with an upright portion that doesn't touch the ground.
  • It likely wouldn't be all that fast but you can probably make an argument that it could move pretty quick in a strait line.

You could also have a tree-dwelling, monkey like creature, I can see an extra set of limbs being great for climbing.

  • All six limbs would need to have gripping digits. Like monkeys these don't have to have opposable thumbs, the can simply hook to latch onto tree branches.
  • If you are going to have thumbs it makes the most sense to develop them on the middle set of arms. Using the top and bottom set for climbing would be the most stable, then the middle arms could manipulate whatever while climbing.
  • I see no inherent difficulty with gait in this setup.

Aquatic creatures also make a lot of sense.

  • Extra fins could make the creature faster and more agile in the water.
  • If you want an aquatic creature with actual hands you probably need the extra set of arms to make them remain decent swimmers.
  • I find this creature less likely than the others to develop digits as survival, especially earlier in their evolution is going to depend on speed and agility in the water...not much need to evolve from that.

I struggle to see extra legs being beneficial in the mountains...but it could probably work, life finds a way to evolve in bizarre and unexpected ways all the time. Nothing immediately comes to mind on this one...the extra legs would be a hazard for a mountain goat as an example.

Predators are interesting with this setup...but complicated. I am immediately reminded of this guy from Avatar super awesome looking, and frankly a bit terrifying. Its actually what I was picturing though probably smaller in my sonic hunter question.

enter image description here

The weird thing about predators...rarely if ever are they the biggest creature.

  • In Africa you have lions and cheetahs, they prey on animals much larger than themselves
  • In North America you have wolves and cougars/bobcats which are quite a bit smaller than most of their prey
  • Bears may be a decent example...but grizzly bears primarily go fishing, they don't really hunt deer or anything, they are not equipped to chase things for long
  • In jungles you have a jaguar, they tend to hunt sloths, tapirs and monkeys. So this may be the best example of a predator that goes after smaller prey.

On a creature that is generally on the ground with a small frame, extra legs seem like they could be more a problem than a benefit...

  • I suppose if you altered the frame a bit you could make it work. If you give the creature four front legs and two back legs I think you could get a gait that works. Plus the extra front legs would allow the predator to grip better when they latch onto their prey.
  • When you specialize your digits to help you grab on and tear open something else it's not super likely you are going to evolve human manual dexterity.
  • Predators are high energy creatures which makes size a liability, being bigger requires consuming more food more often, in lean years, be it from overhunting or a drought...that's not great, so a cool as the six-legged battlecat from avatar is...it doesn't seem real practical.
  • It'd be strange but I can picture six legs where while hunting the middle legs are basically wrapped around the creature's abdomen but it looks creepy and weird in my head.
  • As in avatar I can see six legged creatures working better in a world of larger creatures.
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Ultimately, the preference for 4 or 6 legs within a given environment would come down to the environmental pressures themselves. In its simplest form, the problem is one of agility v. strength.

In an environment where your animal has a survival strategy akin to 'tanking', or making themselves as strongly armoured and stable as possible, 6 legs will win because there are more feet on the ground, and that serves the double purpose of providing a more stable platform and more supportive strength for the armour plating you are going build on your animal. Think animals like Stegosaurus, Ankylosaurus, etc. These were heavy, slow creatures whose survival strategy was based on their ability to withstand attack long enough to give them time to damage or scare off their attackers with their spiked or clubbed tails respectively, and of course their scary roar (we assume).

Such animals would have actually benefited from 6 legs because they could have grown bigger (more legs supporting their weight) but the tradeoff is that they would have been slower. In other words, you're trading speed for toughness.

The extra legs add mass to your body, meaning that they adversely impact the square cube law. That is to say, as you scale up your animal it finds it more difficult to jump and agilely change direction faster because there is more mass by comparison to the relative size of torso for a creature with fewer limbs.

BUT; that's not the real impact of the square cube law for increasing limbs on creatures you want to give speed and agility.

More legs doesn't equal more speed
Adding more legs may increase your torque as a creature, meaning that you can climb steeper mountains or go for longer on the straight, but it simply cannot increase your upper speed. It might increase your acceleration from a standing start in some cases, but that's it. In point of fact, it can actually lower your upper speed because you have to find a gait that combines more feet in an efficient sequence and that's not easy to do.

Torso size can be smaller with fewer limbs
Fewer limbs mean that your torso only has to handle a smaller set of appendage points, so not only do the limbs you have get to be more spread out (and therefore not in the way of each other when running), but the torso can be proportionally smaller meaning that you get the real benefit of the square cube law by decreasing body size generally. Also, the shape of the torso can be squatter, allowing you to fit more organs in a more compact space without disrupting gait. All this leads to a leaner, faster creature suited to the open plains.

Standing
Ever seen a Praying Mantis holding its food with its front appendages? It's not standing but it's doing the closest thing it can to that. Why? Because when man's first ancestors started standing, they only had to tilt one pair of appendages; their legs. In doing so, the spine goes from horizontal to vertical immediately (oversimplification, but true enough for the purposes of this answer) because the hips served as a pivot point.

If your 6 legged creatures want the advantage of height as a vantage point from which to see their enemies approaching, which is largely what proto-man was doing, then you don't have a pivot point because you have 2 sets of hips to contend with. Unless you have a spine with a hinge in it, you're not going to stand upright. Remember, centaurs would have a very complicated spinal column that doesn't really work in the real world because you're essentially attaching an extra spine where the horse's neck normally goes. This is highly unlikely to develop in concert with using the forelegs as hands, and would be a separate and more complicated journey to uprightness than a simple pivot in the hips.

So; the general rule of thumb is;

6 legs - Tankers, slow and cumbersome, but heavily armoured and strong.
4 legs - Faster and agile, more likely to stand upright and develop 'hands' at some point.

As such, your 4 legs are more likely to develop on savannahs or in a world of ambush predators, your 6 legs will develop around large predators that prefer a fight before dinner.

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Evolutionary selection pressure will always favor the solution that has the best cost/performance.

If having 4 legs is good enough to move around, then you will not see any animals with 6 legs.

Having 6 legs has upsides and downsides for animals

Upside:

  • more stability, easier movement on hilly/accidented terrain

  • can survive loss of a leg or two

Downside:

  • movement speed is slower

  • higher energy cost

On Earth the downsides were bigger than the upsides so 4 legs became the favored solution

You could see 6 legged animals with different evolutionary pressures. A planet with high gravity (think 2 or 3g) combined with a very accidented terrain with almost no flat land would probably favor 6 legged animals.

In those conditions, having more stability is worth the extra cost and lower movement speed.

On a planet with even higher crushing gravity, you d probably have animals with no legs. Crawling like snakes or worms would be the favored solution to move around.

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A creature that walks on 4 legs and carries objects with the other 2 legs would need good support if their torso bends upwards.

Either strong muscles and/or bones arranged in a way to support most of the weight. Think about how easily humans pull out their backs when bending over to pick up something.

Such a creature might be better off carrying objects by bending down and under, holding their cargo against their bellies. This would require less effort, but would only make sense if their back or middle legs where for carrying. Otherwise how would they see where they were going?

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  • $\begingroup$ Humans however are mammals which are really poorly designed to be large, mammalian breathing mechanism has stuck them with holes in their torso, meaning all the weight is supported by the spine alone. Dinosaurs on the other hand have nearly rigid torsos becasue the ribs and gastralia go all the way down to the pelvis. that's why you can have things like birds an theropods, that is horizontal bipeds. $\endgroup$ – John Jan 14 at 22:25
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Your question reminded me of the 'hexapoda' (six-legged) dominant life forms on pandora, from Avatar (The movie, not cartoon). Some examples are the six legged horse, the 'dire horse' and the feline-like animal called a 'viper wolf'.

However to answer your question, four is the preferable number of limbs.(A good example of course being primates).Adding more limbs would just require more brain-power to control them, and would require a major change in configuration that would mean that it would have to have some benefit.

  • While we don't have dire horses or 'viper wolves' on earth, we do know one species that have six limbs- insects. When a creature runs on two or four legs, only one or two feet remain in contact with the ground at any point in time. When an insect runs on a surface, three limbs usually remain in contact with the ground at all times. For insects like a fly, for example, this is an advantage because it means it has greater stability when clinging to a vertical surface or upside-down. That's not an advantage that other animals need walking upright on a flat surface.
  • In many ways, six legs would hinder normal creatures. Insects on Earth that have six legs have them splayed out from the body so they don't get in each other's way. If your life form is anything like Avatar's, then all the creatures would have their six legs in line with their bodies, which would cause them to bump into or get tangled up together easily. As shown with our own history and science, four legs seems to be the best recipe for land-dwelling animals. Unless your creatures climb vertically etc, then four legs would be much more beneficial.

  • Furthermore, creatures with more than four limbs have not evolved in land species. In fact the trend is to 'simplify'- as with snakes,
    which lost their limbs- and horses, which instead of having arms and hands, have hooves. Six, eight and even leggier arthropods (insects, arachnids and crustaceans) have, of course, been extraordinarily
    successful. But the fact that these creatures have exoskeletons
    rather than bones inside their bodies limits their size.

  • Large hexapoda animals would need Large exoskeletons to support
    internal body tissues etc. However, it would be prohibitively
    cumbersome and heavy, and would demand lots of energy to move and
    grow. Because intelligence is strongly linked to how big, intricate
    and energy-consuming an animal's brain is, that sentience is unlikely to evolve in hexapoda like animals.

Edit: I concentrated on listing the disadvantages of having six legs, rather than directly comparing it with four legs.

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    $\begingroup$ "would require a major change in configuration that would mean that it would have to have some benefit" On the flip side, going from six to four limbs would also be a major change. Evolution usually ends up at local, not necessarily global, maxima. For a six-legged creature to evolve toward becoming a four-legged creature, each intermediary form would need to, at best, not confer a disadvantage. So the same argument could be made both ways. Also, in the question, I specifically noted that answers could ignore the fact that some life in this world has a different number of legs from 4 or 6. $\endgroup$ – a CVn Jan 14 at 17:59
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Given the example of the development of the scorpion, it started out as a segmented worm with a pair of legs/swimming paddles per segment. These legs later developed into a variety of appendages, such as pedipalps, mandibles, pincers, and land traversing legs.

Beetles similarly developed palps, maxilla, mandibles, wing case lifting appendages, a set of four wings or wing counter weights, and six legs. If you consider the specialization of legs into structures as mentally nimble as a scorpion's pincers or a grasshopper's wings. I don't see any inhibition for a six legged terrestrial or possibly arboreal animal in development of manipulative appendages.

If the structure of any creature has a survival advantage the species tends to retain the structure. The concept that it is too mentally taxing to control large numbers of limbs is disproved by looking at the large number of multilegged creatures with rudimentary brains. A shrimp has four sets of manipulative maxillapeds, two sets of periopeds (walking legs) and five sets of pleopods (swimming legs) and two sets of uropods (fins) in its tail. With its undeveloped microscopic brain it can coordinate all twenty six of them to work in unison to locomote, maneuver and manipulate things in its environment.

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If you use all your limbs for locomotion, then 4 makes sense because you can lift one while maintaining balance on the other three. If you only had 3 legs, you would have to maintain balance on two of them whenever you moved, and if you can do that easily then you don't need more than two legs for locomotion. If you had more than 4 legs, then you don't gain anything with the extras.

That said, it benefits bugs to have more legs because of their other functions, such as climbing/hanging/landing on ceilings, quickly navigating/braiding complex webs, or being able to lose segments without dying.

So, for a large creature to justify having more than 4 limbs, it would need to have some needful task or danger which merits the extra limbs. Some example ideas:

  • The planet has creatures who are adept at attacking from behind (for some reason), and so the other creatures evolved an array of sensitive hairs and an extra set of limbs on their backside to detect and deflect such attacks
  • The planet is covered in pools of fluid with high surface tension, so having more legs disperses the creature's weight enough to let them run across those pools with relative ease.
  • The creatures fly and need to be able to land on very complex/porous/webbed terrain.
  • The creatures primary source of food tends to hang off the edges of cliffs, so they need to be able to support themselves by gripping the cliff edge and food stalk, without putting too much pressure on the stalk itself.

I'm sure there are endless good reasons for having more than 4 legs, but given the ecosystem and environment on planet earth, most of our creatures have exactly the number of legs that they need; more would be wasteful and fewer would make survival difficult.

Hope this helps!

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So from my understanding of the situation, the early dominant species to walk on land in this world were six legged and they went on to evolve into many other animals, some of those descendants went on to evolve into four legged animals with two arm like appendages.

Why would animals have evolved this way though? Most obvious answer would be speed. You don't need six legs to gallop.

Hopping would be an alternative (just think of fleas or other insects of that ilk), but that becomes problematic as the animals get bigger and probably not great in certain terrains (un-even ground or densely wooded areas).

I envisage that at some point a common ancestor of your six limbed (four legged, two armed) animals developed running on four legs and picked its fore limbs up off the ground to do this. Natural selection kicks in and the two pairs of back legs get further apart and the front legs become smaller. Animals which survive by pure strength/stability may stay on six legs.

Some animals may lose the use of the fore arms altogether if they find no evolutionary need for them.

But other animals could find different uses for those fore limbs like pulling plants up to get at the roots, latching onto prey, pick fruits from trees, mating rituals/displays or building things (nests, burrows, dams, etc.). These are the sort of activities that may spur evolution in the direction of fingers/thumbs and fine manipulation.

Of the animals that did find a use for their fore limbs, they could end up like your desired state animal of being four legged, with the ability to carry and manipulate things in a way similar to humans.

Only thing I would say, is that I can't see an obvious evolutionary path to have animals with an upright front half (centaur like). More likely that the animals limb joints (shoulders, hips, mid-hips?) would line up as opposed to forming an angle. As mentioned in another answer, a centaur would have great difficulty in picking something up off the floor. Although it's head would need to be above the shoulders so the arm like limbs don't interfere with peripheral vision.

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There is no real disadvantage other than the material cost you get with any limb, but keep in mind tissue needs a use to be kept over prolonged evolutionary time so make sure your "missing link" species also has a use for them. The main reason modern tetrapods only have four is they evolved from fish that also had four limbs.

there is one notable exception for an animal with an upright posture, 4 limbs has a distinct advantage because six limbs start to get in each others way during the stride, so either you need a much longer (and thus more massive) creature or you need to be slow moving.

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One disadvantage that I can think of is they would not be able to run fast. Take for example a Jaguar or a Deer. If you add two more limbs they would get in the way of running.

When a Jaguar runs it leaps with the front legs and catches up with the hind limbs where hind limbs cross the front legs/limbs. If you add two more in the middle they would get in the way.

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  • $\begingroup$ Hi saurav, welcome to Worldbuilding! We hope you enjoy your stay. Check the help center and take the tour if you haven't already! Currently, you talk about the running speed of such an animal. Could you elaborate on how the limbs would get in the way of running? $\endgroup$ – JavaScriptCoder Jan 14 at 19:41
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    $\begingroup$ When a Jaguar runs it leaps with the front legs and catches up with the hind limbs where hind limbs cross the front legs/limbs. If you add two more in the middle they would get in the way. $\endgroup$ – saurav kumar Jan 14 at 19:47
  • $\begingroup$ ohh and also Hello there !! :) $\endgroup$ – saurav kumar Jan 14 at 19:48
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    $\begingroup$ @sauravkumar Greetings and welcome! Would you mind editing your comment response into you actual answer? Just keep in mind: the whole point of Stack Exchange is to provide the best answers possible. We shouldn't force people to read through all the comments just to piece together a coherent response. $\endgroup$ – elemtilas Jan 14 at 20:05

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