Arthropods have 6 or more limbs and arthropods with 6 limbs appear to move faster than arthropods with 8 limbs so I wonder whether this might have something to do with fast and efficient locomotion. But, this is just a guess. I wonder what the official explanation is, if it exists.

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    Basically because mammals and arthropods evolved separately. If we evolved from a four-legged, fish-like ancestor, then mammals never got to choose how to evolve more efficiently (if it is). – anongoodnurse Oct 1 '14 at 0:14
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    @anongoodnurse What does "efficiency" have to do with anything? If the theory that everything evolved from single-celled organisms is correct, then that argument holds little water. – b1nary.atr0phy Oct 1 '14 at 9:36
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    @b1nary.atr0phy - how so? – anongoodnurse Oct 1 '14 at 13:25
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    Are prehensile tails not considered limbs in this case? Common definitions seem to include them as limbs, giving many mammals 5 limbs instead of 4. – tpg2114 Oct 1 '14 at 16:20
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    I think that if mammals had 6 limbs, you could ask why don't mammals have 8 limbs. Or if they had 8 limbs, you could ask why wouldn't they have 6 limbs. Evolution had to act a certain way, and it worked in the way so that present day mammals don't have more than 4 limbs. – JonHerman Oct 2 '14 at 1:17
up vote 78 down vote accepted

Number of legs in terrestrial vertebrates

Not only do mammals have four legs but actually all terrestrial vertebrates (which include mammals) have four legs. There are slight exceptions though as some lineages have lost their legs. Typically snakes have no legs anymore. Apesteguia and Zaher (2006) discuss the evolution of snakes legs reduction and report a fossil of snakes with a robust sacrum. Cetaecea (whales and friends) have lost their hind legs but we can still spot them on the skeleton. See for example the orca (killer whale, easily recognizable to its teeths) on the picture below. Pay attention to the small bones below its vertebral column at the level on the left side of the picture.

I also want to draw attention to the importance of the definition of legs. I guess that we would call something a pair of legs if it is constructed using a similar developmental pathway than current existing legs. If we are using some broader definition, then a prehensile tail as found in some new world monkeys for example could be considered as a leg (but only a single leg, not a pair of legs obviously). A list of animals having a prehensile tail can be found here (wikipedia).

Did you say Natural Selection?

I think (might be wrong) that you have too selectionist a view of evolution. What I mean is that you are wondering why mammals have four legs and you're looking for an explanation of the kind "because mammal have this kind of need of locomotion and for this purpose four is the most optimal number of legs". Consider the following sentence: "If there is a need, natural selection will find a way!". This sentence is wrong! Evolution is not that easy. This falsy view of evolution is sometimes reffered as panselectionist.

The reality is that it is not easy to evolve such a developmental pathway as drastic as having an extra pair of legs that is well integrated into the body of the carrier of this new trait. Such individual would need a brain, a nerve code, a heart and some other features that are adapted to have extra legs. Also, assuming such a thing came to existence it is rather complicated to imagine how it could be selected for. To go slightly further, you have to realize that there are many stochastic processes in evolution (including mutation and random variation in reproductive success) and an organism is a complex machinery and is not necessarily easily transformable to some other form that would be more efficient (have higher reproductive success). Often going from one form to another may involves a "valley crossing" meaning that if several mutations are needed, intermediate forms may have low reproductive success and therefore a high amount of genetic drift (stochasticity in reproductive success) to cross such valley of low reproductive success. See shifting balance theory. Finally, even if there is selection for another trait, it may take time for the mean trait in the population to shift especially if there is only little genetic variance. A complete discussion on why the sentence "If there is a need, natural selection will find a way!" is wrong would fill up a whole book.

Gould (1979) is a classical article on the subject and is very easy to read even for a lay person.

Why 4 legs?

Terrestrial vertebrates have four legs because they evolved from a fish ancestor that had four members that were not too far from actual legs (members that could "easily" evolve into legs). This is what we call a phylogenetic signal. The explanation is as simple and basic as that. You can have a look to the diversity of terrestrial vertebrates here (click on the branches).

Number of legs in invertebrates

Arthropda (Spiders (and other chelicerata), insects (and other hexapods), crustaceans (crabs, shrimps…) and Myriapoda (millipedes) and Trilobite as well)) evolved from a common ancestor who had a highly segmented body. From this ancestor many group have fused some segments. In these taxa, each pair of legs is attached to a particular segment (I don't think the segments are still visible in spiders today). In insects for example, all 6 legs are attached to the thorax but to 3 different segments of the thorax, the pro- meso and meta- thorax (see below).

As a sidenote, it is interesting to know that the wings in insects did not evolve from the legs (as it is the case in birds and bats). There are two competing hypotheses for the origin of insect wings. Wings either developed from gills or from sclerite (chitine plate, the hard part of the insect). When insect first wings, they actually evolved three pairs of wings (one on each segment of the thorax). At least one pair has then been lost in all modern species. In the diptera a second pair of wings has been lost and are replaced by halteres, particularly easy to spot in craneflies (see below picture). In millipedes the link between segmentation and legs is even more obvious (see picture below). You can have a look to the diversity of Arthropda here (click on the branches).


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Update 1

Asking how likely it is for a given population to evolve a given trait is extremely hard to answer. There are two main issues: 1) a definition of the question issue and 2) a knowledge issue. When asking for a probability one always needs to state the a priori knowledge. If everything is known a priori, then there is nothing stochastic (outside quantum physics). So to answer the question one has to decide what we take for granted and what we don't. The second issue is a knowledge issue. We are far from having enough knowledge in protein biophysics (and many other fields) to answer that question. There are so many parameters to take into account. I would expect that creating a third pair of legs would need major changes and therefore one mutation will never be enough in order to develop a third pair of legs. But, no I cannot cite you any reference for this, I am just guessing!

Following the wings example in insects. Insects have had three pairs of wings. While some mutation(s) prevented the expression of the third (the first actually) pair many of the genetic information for this third pair remain in the genotype of insects as they still use it for the two other pairs. Taking advantage of that, Membracidae (treehoppers) developed some features using similar biochemical pathway than the one use to develop wings. Those structures are used as protection or batesian mimicry.

Update 2

Let's imagine that an extremely unlikely series of mutations occur that create some rodent with 6-legs. Let's imagine this rodent with six legs has a larger heart in order to pump blood to these extra legs and it has a brain that is adapted to using six legs and some changes in its nerve cord so that it can control its 3rd pair of legs. Will this rodent have higher reproductive success than other individuals in the population? Well… let's imagine that with its six legs, it can run faster or whatever and has a very high fitness. How would the offspring of a six-leg mother (or father) and a four-leg father (or mother) look like? Will it be able to reproduce? See the issue is that it is hard for such trait to come to existence because 1) it needs many steps (mutations) and 2) it is hard to imagine how it could be selected for. For those reasons, there exist no vertebrates with 6 fully functional legs.

Well, let's assume it does and in consequence after 200 generations or so, the whole population is only made of 6-legged individuals. Maybe the species got extinct then and no fossil record has ever been found. This is possible. It is not because something has existed that we necessarily find something in the fossil record.

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    @anongoodnurse: Depending on your definitions, conjoined twins might provide a counterexample to your assertion. And of course, some mammals have evolved wings, so it seems reasonable to ask why our particular species did not. – Nate Eldredge Oct 1 '14 at 16:05
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    @NateEldredge - you're not reading my comments properly. Conjoined twins are a developmental, not genetic, defect, and bat wings are not from a third pair of limbs. – anongoodnurse Oct 1 '14 at 16:25
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    Nate I encourage you to lookup pictures of bat skeletons. Their skeleton looks like a person with really big hands, because that's basically what bat wings ARE: really long fingers with skin stretched between them. In fact, bats are one of the best illustrations of how all mammals evolved from a common ancestor and thus have basically the same body structure (ie. the answer to this question). – jhocking Nov 24 '14 at 13:35
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    @anongoodnurse a mutation in the Hox genes can cause formation of additional limb buds – WYSIWYG May 14 '15 at 18:56
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    I think having an additional limb isn't that rare in mammals. Having one which is usable to a degree that it increases fitness (i.e. chances of reproduction) is apparently much rarer. People who solder or cook for a living would love to have another arm... @anongoodnurse: Beauty is in the eye of the beholder; what some call a "defect in embryonic development" may be an enhancement to others. And don't tell me embryonic development is not dependent on the genes... – Peter A. Schneider Sep 8 '16 at 14:39

I think I might interpret your question as asking, not just why don't mammals have more than four limbs, but why arthropods have more variety. Insects have six, but others have eight, ten, or more.

Partly there are just many more species of arthropods. 80% of animal species are some sort of arthropod, and some lineages of arthropods are distantly related to each other because arthropods as a group have existed since the Cambrian period. With so many more species, it's no surprise that they've managed to tack on some extra variety.

Although it might seem that insects and spiders are more closely related than, say, humans and fish, the reverse is actually true. In fact, insects split off from other arthropods at approximately the same time that fish were diversifying into the three main lineages that would eventually become cartilaginous fish (sharks & rays), ray-finned ("regular") fish, and lobe-finned fish (coelacanths and lungfish). All of these things happened at approximately the same time, geologically speaking, at around 400 million years ago. Some ancient lungfish crawled up onto land a short while after, giving rise to land animals. So in light of this, it's not really surprising that arthropods would have so much diversity in body plan - you don't look much like a leafy sea dragon (I hope).

But perhaps another way of looking at it is in the difficulty involved in adapting the body plans of these different types of animals. Arthropods have segmented bodies. In some arthropods, especially the myriapoda (centipedes, millipedes and so on), there's a pretty straightforward arrangement of one segment to one pair of legs (excluding the head). Vertebrates have a segmented body plan too, but our segments aren't quite so well-separated. In vertebrates, a "segment" is approximately equal to one vertebra. So since vertebrates aren't in the habit of attaching a pair of legs to each vertebra, it's harder to grow more legs. Instead, vertebrates have internal skeletons, with the familiar hips and shoulders in land animals. Vertebrates do have good luck adapting the internal structure when they need to. Most use the traditional four-leg layout, but humans, birds, whales, and snakes have all made pretty significant changes to it. But internally, it's still the familiar hips and shoulders - even snakes still have the necessary bones. Given another few hundred million years, they might disappear entirely, but it seems like the basic bone structure is tough to change. Who knows, in a hundred million years, the far future descendants of snakes might end up with legs again, and grow six or eight instead of four!

So, overall, I'd say that the two main reasons are that the arthropod body plan is just a little more flexible due to its higher degree of modularity, and that different kinds of arthropods are actually more distantly related than you might expect.

The answer by Remi.B is excellent, I'll just attempt an explanation by way of gene networks:

In genetics we see new genes "linking" to the older genome by regulation pathways and by being "fit" only in the context of the existing genome. This has the effect of making the older genes indispensable. Change them and you rupture the whole mesh. If you want to increase the number of limbs, you'd have to take that up with the HOX genes, that control the basic body plan. These are awfully ancient, originating in the common ancestor of flies and humans [1]. So as a rule of thumb, these genes will also be awfully hard to mutate without ruining everything.

Another line of reasoning is that the HOX genes not only appeared early in evolution, but they are also expressed early in the development of the embryo. When anything that goes on in development depends on what has preceded it, mutations in HOX genes has a bigger chance of ruining something than mutations that are only expressed late in development. I'm going out on a ledge here, but consider that the color of hair, skin and eyes of a human baby are often not fully expressed until months after its birth and consider also the range of harmless variations in these colors. These superficial colors don't affect anything else, so the corresponding genes are free to mutate into a great variety.


Here is a more morphologic, less genetic answer:

According to Sansom 2013, the 2 sets of paired appandages (shoulder and pelvic) was set in stone when agnathans transitioned into gnathostomes (ie. when the first vertebrate organisms began to evolve jaws, an anatomic change that allows for classification of different stages of history found in the fossil record). They found variations where an agnathan had a paired set of anal fins (sort of the 3rd set of paired appendage, since anal fins are typically unpaired).

Now, moving forward millions of years, the Sarcopterygii (lobe fin fish) evolved into the tetrapods (4 limbed land vertebrate including amphibians, reptiles, birds, and mammals), this pattern (2 sets of paired limbs) became immutable, though much less fundamental variation throughout the phylogenesis of the various tetrapods.

Sansom RS,Gabbott SE, Purnell MA. 2013 Unusual anal fin in a Devonian jawless vertebrate reveals complex origins of paired appendages. Biol Lett 9: 20130002.

Don't forget the laws of physics...

Yes, tetrapods (mammals, birds and herptiles), may have initially inherited genes for two pairs of limbs from their fish ancestors, but why do fish themselves have limited numbers of fins, rather than sixteen or 400?

Moreover, some tetrapods have lost one or even both pairs of limbs.

To put it in perspective, how many invertebrates can you think of that have just two pairs of legs? If vertebrates can lose limbs, so can invertebrates. Yet Nature seems to favor greater numbers of limbs among invertebrates.

I don't know exactly why, but I suspect it could be something as simple as physics. Imagine an elephant with six massive, pillar-like legs. The animal would be significantly heavier and therefore slower. Plus, it would need more nutrition to support the extra limbs.

So I would argue that the answer is a combination of genetics and "physics," in the broad sense of the term.

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    Please could you add some supporting scientific material that reinforces your answer and allows further reading. – ELL Mar 8 '16 at 7:25
  • "...something as simple as physics. Imagine an elephant with six massive, pillar-like legs. The animal would be significantly heavier and therefore slower. Plus, it would need more nutrition to support the extra limbs." I do not know if you know exactly what the word "physics" entails. There are so many holes in the argument that a cannon ball can shoot through without grazing a hair! – Hans Sep 26 '16 at 21:58
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    For a starter, the elephant can have six thinner legs with the same total cross sectional area as before and thus the same, NOT more, total mass and thus about the same nutrient to support them as with four limbs. Now the only thing that may require a bit more energy is the total surface area of the legs is $\frac{\sqrt{3}}{2}$ times as the four legs and increase the surface heat loss and may require more energy to compensate for the heat loss but also good for cooling down. – Hans Sep 26 '16 at 21:58

protected by ELL Dec 10 '14 at 1:15

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