Darwin right again: the inner ears of sloths are highly variable

Darwin said a lot of good stuff in The Origin. One of his less-recognized achievements was to explain not just the presence of rudimentary (“vestigial”) features as evidence for evolution, but also to recognize that such features, insofar as they’re not essential to the individual, would be more free to vary than would features critical for survival and reproduction. Here’s his discussion from Chapter 13: “Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Organs.” (I’ve put the sentence most important for this post in bold.)

On my view of descent with modification, the origin of rudimentary organs is simple. We have plenty of cases of rudimentary organs in our domestic productions, as the stump of a tail in tailless breeds, the vestige of an ear in earless breeds, — the reappearance of minute dangling horns in hornless breeds of cattle, more especially, according to Youatt, in young animals, and the state of the whole flower in the cauliflower. We often see rudiments of various parts in monsters. But I doubt whether any of these cases throw light on the origin of rudimentary organs in a state of nature, further than by showing that rudiments can be produced; for I doubt whether species under nature ever undergo abrupt changes. I believe that disuse has been the main agency; that it has led in successive generations to the gradual reduction of various organs, until they have become rudimentary, as in the case of the eyes of animals inhabiting dark caverns, and of the wings of birds inhabiting oceanic islands, which have seldom been forced to take flight, and have ultimately lost the power of flying. Again, an organ useful under certain conditions, might become injurious under others, as with the wings of beetles living on small and exposed islands; and in this case natural selection would continue slowly to reduce the organ, until it was rendered harmless and rudimentary.

Any change in function, which can be effected by insensibly small steps, is within the power of natural selection; so that an organ rendered, during changed habits of life, useless or injurious for one purpose, might easily be modified and used for another purpose. Or an organ might be retained for one alone of its former functions. An organ, when rendered useless, may well be variable, for its variations cannot be checked by natural selection.

Note the important point—often neglected by creationists—that vestigial organs can be functional (“Any change in function, which can be effected by insensibly small steps, is within the power of natural selection; so that an organ rendered, during changed habits of life, useless or injurious for one purpose, might easily be modified and used for another purpose.”).  Thus the wings of penguins, no longer used for flying, are useful in helping them swim, and at the same time testify to penguins’ origin from flying birds. They are useful vestigial traits.

There are are least three reasons why a now-useless feature can be eliminated. First, mutations affecting it are no longer deleterious, so it would degenerate over time. Second, the feature could be injured and, if no longer useful, would be eliminated by natural selection because its injury would reduce fitness (this may be the reason why the eyes of cave animals gradually disappear). Finally, the metabolic energy that goes into producing a useless feature could be diverted to other features that are still useful (producing a useless organ like an eye has physiological “costs,” and maybe you could divert the metabolic resources involved in making it into other features, like the brain).

From all of these, but particularly the first, you can derive Darwin’s conclusion why useless features should be more variable than essential ones. In the technical jargon of evolutionists, those features are no longer under “stabilizing selection”, the kind of natural selection that eliminates deviations from features that are at a local optimum for being fit.  Usually mutations causing deviations from such a feature are selected against, but may no longer be deleterious if the feature is no longer useful.  Mutations can thus accumulate willy-nilly, and the feature becomes more variable.

The variability of vestigial organs is well known. The classic example is that of wisdom teeth, which are highly variable in both presence (Tasmanian aboriginals don’t have them; indigenous Mexicans all have them), and degree of expression, as many of you know.  Some people can’t wiggle their ears because their ear muscles aren’t functional (we no longer need them since we can localize sound by moving our heads), while others, like me, can wiggle them freely.  The palmaris longus muscle has been reduced to a small tendon in our forearm; it’s absent in 14% of people and, being pretty useless, is often used for tendon grafts in the wrist.

Further support for Darwin’s notion of increased variation in less-useful characters comes from a new study of variation in the inner ear of the three-toed sloth (Bradypus variegatus). A  paper in the Proceedings of the Royal Society by Billet et al. (download free; reference below) shows that several features of the inner ear, a part of the body crucial for maintaining balance, are much more variable in the sloth than in its relatives.  This is expected if sloths, being notoriously slow, don’t have a critical need to maintain balance.  The authors were particularly concerned with the semicircular canals (SC) of the animals, which are really crucial for keeping balance. There are three semicircular canals, each of which contains fluid and small hairs that detect the motion of the fluid. In that way the animal is able to sense its orientation.

As Billet et al. note:

Accordingly, one would expect to find among sloths a substantial degree of morphological variation in their SC, because significant travelling distances, speed and agility are not part of their locomotor repertoire. In this context, sloths constitute an ideal case to address Darwin’s hypothesis.

Well, sloths still have to hang in trees and maintain balance when they defecate (they do so once a week, climbing all the way down from their trees to do their business at the base of the tree before climbing back up, a behavior that’s not really understood), but maybe they don’t need a perfectly-formed inner ear to do that, so variations caused by mutation or developmental accidents might not be terribly injurious.

To test “Darwin’s hypothesis,” the authors used high-resolution computer tomography to examine and make 3-D reconstructions of the inner ears of individual three-toed sloths as well as of red squirrels (Sciurus vulgaris), European moles (Talpa europaea), the collared anteater (Tamandua tetradactyla), and the nine-banded armadillo (Dasypus novemcinctus). The anteater and armadillo are members of the Xenarthra, a suborder of New World mammals that include the sloths, and thus these species serve as “controls” for degree of variation in related species.

Billet et. al measured 12 features of each inner ear, and, since variation might be correlated with size, expressed variation as the variation among individuals in the ratios of one feature to another.  Here’s the inner ear in situ of a three-toed sloth skull, and its reconstruction from the computer scan (“ASC,” “PSC” and “LSC” are the three semicircular canals, also called “horizontal,” “superior,” and “posterior”):

I’ll show the numerical values of variation in a second, but this figure from the paper shows the higher variation in sloths just by inspection of the degree of variation among individuals of four species. The sloth is clearly more variable than the anteater, armadillo, and squirrel. As you might expect, the squirrels show almost no variation:

Here are the data, with the height of the bar showing the degree of variability of ratios of inner-ear features. The red bars are the sloths, green bars are anteaters, purple bars are armadillos, yellow bars are squirrels, and blue bars are moles. In all cases the variation is highest in sloths, and in many cases it’s much higher than the other species (in nearly all cases the difference between sloths and other beasts is statistically significant):

The authors conclude that they’ve verified Darwin’s hypothesis and advance this reason for their observation:

Therefore, we hypothesize that relaxed selective pressure on the SC has permitted the persistence of the observed variants in three-toed sloths. The level of genetic and/or developmental constraints on the production of these variants might also depart from that of other placentals. . . The most plausible reason for such a released selective pressure on their SC morphology lies in their reduced activity pattern. Such high variation of the SC has not been detected in faster-moving xenarthrans, nor has it been documented in other fast-moving mammals.

It’s known that animals that are agile and move fast have relatively larger semicircular canals, so I find this conclusion plausible.  What remains is a more definitive test of whether variation in the inner ears of sloths really does matter less than such variation in other species. I’m not sure how one would test that, for you can’t just go in and just manipulate the inner ear. Perhaps you could put sloths in jet planes, subject them to various sorts of vertigo, and see if they do better than other animals!

Another possibility is that sloths are simply more variable than related mammals in many features, and so the ear variation isn’t that special.  Maybe, for instance, the sloths measured came from genetically isolated subgroups, and the variation we see is just variation among subspecies or ecotypes.  The authors argue pretty convincingly that this isn’t true, but at least two other studies show substantial variation in other skeletal features of sloths that aren’t related to balance, and in the timing of when their cranial sutures (the fissures in the skull) close.  So we need to look at variation in many morphological traits of sloths.  Given the funding difficulties of this type of science, I doubt we’ll see such work in the near future.

One thing I found amusing was the blurb for the short HuffPo piece on this research:

It sort of suggests that Darwin’s theory in general was under question, rather than just this one small prediction that Darwin made. But even that prediction has already been verified in characters like wisdom teeth and human muscles, so the present study doesn’t really add much to substantiating that idea.  What the work does suggest, though, is that it’s less important for sloths than for other animals to maintain their balance, and that’s cool in itself.

Finally, here’s a video of a three-toed sloth (and its faster two-toed relative, which isn’t all that closely related):

“Sloth” of course, is an adjective that applies to lazy humans. That reminds me of a joke: A city boy goes to the country and sees a bunch of hogs slopping noisily and messily at their trough. “No wonder they call them pigs!”, he exclaims.

I’ll be here all week, folks.


Billet, G., L. Hautier, R. J. Asher, C. Schwarz, N. Crumpton, T. Martin, and I. Ruf. 2012. High morphological variation of vestibular system accompanies slow and infrequent locomotion in three-toed sloths. 10.1098/rspb.2012.1212. Proceedings of the Royal Society B: Biological Sciences


  1. gbjames
    Posted August 5, 2012 at 8:22 am | Permalink

    Very interesting post with lots of little tidbits I didn’t know. I had no idea that Tasmanian aboriginals don’t have wisdom teeth!

    • Filipe
      Posted August 5, 2012 at 3:08 pm | Permalink

      Didn’t is probably more adequate. Modern aborigines probably have them, since they have a fair share of European ancestry. Tasmanian aborigines were subject to genocide in the early 19th century leading to near eradication (they were even declared extinct in 1876).

    • Anaxyrus terrestris
      Posted August 5, 2012 at 7:04 pm | Permalink

      Actually, Jerry got this one backwards. If you read the linked wikipedia source, or the original abstract from which the tidbit was gleaned, you see that it is agenesis (lack of wisdom tooth formation) that is 0% in Tasmanians (they all had wisdom teeth) and 100% in the sample of aboriginal Mexicans (they all lacked wisdom teeth).

  2. danarel
    Posted August 5, 2012 at 8:55 am | Permalink

    creationists: “satan gave sloths ears to test your faith”

  3. BilBy
    Posted August 5, 2012 at 9:04 am | Permalink

    Never knew about the wisdom teeth, never knew about the palmaris longus muscle – thank you!

  4. ManOutOfTime
    Posted August 5, 2012 at 9:54 am | Permalink

    Regarding the “Darwin Proved Right” headline, I find it funny that a news and infotainment outlets impose style guides on their writers about things that don’t matter to most of us – the use of serial commas, referring to people as Mr. and Ms rather than just their last names, etc – but when it comes to a scientific subject, they without fail use ignorant or out-dated tropes. Dude, there is no “missing link.” Darwin, or Einstein for that matter, are not being challenged or “proven right” by each additional paper, as if we have not yet achieved a preponderance of the evidence.

    The Sideboob Gazette may not have such standards at all, not being a top-down organization. On the other hand, being an accommodationist organ, you can’t help but wonder if the particular phrasing isn’t at least a symptom of that culture and willfully, woo-fully ignorant worldview.

  5. Posted August 5, 2012 at 10:08 am | Permalink

    I can see the Onion article now:

    Darwin was right, the frontal lobes of the religious are highly variable.

  6. Ougaseon
    Posted August 5, 2012 at 10:11 am | Permalink

    Jerry, how well supported are hypotheses about the metabolic costs of maintaining rudimentary organs as a factor in natural selection? It has often seemed to me that the other two explanations you mention are both much more convincing a priori and easier to actually test.

  7. Adam Felton
    Posted August 5, 2012 at 10:13 am | Permalink

    Ditto! What a great post!

    • jimroberts
      Posted August 5, 2012 at 10:32 am | Permalink


  8. Scott near Berkeley
    Posted August 5, 2012 at 12:44 pm | Permalink

    Jerry, great post, quite informative, plenty of trivia (e.g. wisdom teeth) and wonderful examples to demonstrate scientific examination of the real world, at its best.

  9. marksolock
    Posted August 5, 2012 at 1:01 pm | Permalink

    Reblogged this on Mark Solock Blog.

  10. MNb
    Posted August 5, 2012 at 1:44 pm | Permalink

    Great post indeed, especially as it shows how cute sloths are.

  11. Mary - Canada
    Posted August 5, 2012 at 2:34 pm | Permalink

    I remember the first time I saw a sloth on tv (long time ago)….was the most amazing animal I had ever seen. Thanks for the post.

  12. zuzanafan
    Posted August 5, 2012 at 5:38 pm | Permalink

    I always enjoy these interesting posts about evolution. It’s always fun to learn more about how the theory of evolution makes testable predictions. Every time I read a post like this I think of my creationist upbringing, but as time goes by I start to just appreciate learning about the world and not letting creationist retardation come into my thoughts.

  13. Notagod
    Posted August 5, 2012 at 7:18 pm | Permalink

    shows that several features of the inner ear, a part of the body crucial for maintaining balance, are much more variable in the sloth than in its relatives.

    Could it be that the three-toed sloth would benefit from a better balancing mechanism but the sensitivity of the components weren’t up to the task of providing the sloth any benefit due to its very slow movements? That is, maybe the evolution of the balancing mechanism’s sensitivity wasn’t able to keep up with the selection pressure for slower sloth movement? Maybe if the variation happens upon a combination that provides for a working balancing mechanism the variation will stabilize?

    • Launcher
      Posted August 6, 2012 at 5:21 pm | Permalink

      > That is, maybe the evolution of the balancing mechanism’s sensitivity wasn’t able to keep up with the selection pressure for slower sloth movement?

      I think you’re right, and I refer you to my comment below. The fluid/hair cell coupling in the semicircular canals is not effective at very low frequencies (long time constants) of rotation. There is a “central integrator” in the mammalian brainstem, neural circuitry that effectively extends the time constant of the canal signal, but only so much. Instead, I speculate that the sloth relies on alternative sensory cues for balance and orientation in space, especially vision and the otolith system, which encodes translation and tilt.

      There may be a reverse analogy here with people that suffer from bilateral vestibular deficits. They cope with the loss of vestibular input by slowing down their movement behaviors and relying more on visual information.

      • Notagod
        Posted August 7, 2012 at 7:14 am | Permalink

        Thanks for your explanations Launcher. I wasn’t sure if the sensitivity of the hair movement in the fluid could have a limit within the typical speed of the sloth or not. Its nice to know a little more about how it all fits together.

        • Launcher
          Posted August 10, 2012 at 4:37 pm | Permalink

          My pleasure. I suppose that the viscosity of the canal fluid (endolymph) or the size + mass of the hair cells COULD have adapted to make the semicircular canals more useful to the slowly moving sloth. But it seems this was not to be, so the canals may be on their way to being truly vestigial on this evolutionary branch.

  14. Marella
    Posted August 5, 2012 at 7:40 pm | Permalink

    OMG, I don’t seem to have a Palmarus longus! I always knew I was highly evolved. 😉

    It is amazing how little Darwin was wrong about, especially considering how much work he did in so many fields.

  15. Kevin
    Posted August 5, 2012 at 8:23 pm | Permalink

    “Note the important point—often neglected by creationists—that vestigial organs can be functional”

    I thought it was evolutionists who liked to point to “non-functional” organs as evidence against intelligent design – only to later discover a function.

  16. Posted August 5, 2012 at 10:48 pm | Permalink

    Kind of reminds me, in principle, of inertial navigation systems I used to check out as a QA guy at Autonetics in the 60’s. The stable platforms within the system contained gyros for stabilization, and to maintain spatial orientation.

    Key to spatial location were the three accelerometers, aligned with the x y & Z axis. One inspection point was to verify that the quadrature axis orientations were within spec. When launched, the missile would interpret the accelerometer outputs as changes in its spatial orientation, and with an offsetting voltage to the platforms X and Y axis motors to compensate for earthly rotation.

    While true that the sloth appears to not require as keen a sense of orientation, and thus the sensor tubules were allowed to degenerate. Birds on the other hand have a strong need of that function, in particular with a need to maintain stable flight at times when no horizon is visible.

    I know firsthand, remembering my first takeoff from Falcon Field, Fullerton CA on a smoggy day, and where limited VFR rules were place, requiring normal VFR upon reaching 5000 feet. But at 1500 feet when the ground disappeared from view, disorientation immediately set in. Fortunately, my instructor was there and who pointed my attention to the artificial horizon bar for reference and voilà, the panic subsided.

    So do birds have better developed vestibular system than humans? According to an interesting doctoral thesis, they do. In particular, “raptors, including hawks, falcons and owls exhibit agile behavior” and “volant birds as a group [the above] have larger canals, relative to body mass than bipedal dinosaurs.” And of course, humans.

    Somewhat similarly, the dogfish which has large pectoral fins, depends heavily upon the SC’s. Damage to the VIII nerve or sequestering one of the SC’s has been shown experimentally to cause the fish to roll along its longitudinal axis.

    Thinking back about me at 1500 feet over Falcon Field and with no horizon reference, a slight movement of my pectoral fins, correction ailerons , would cause not just a roll, but a bank, increasing a seat-of-the-pants increase, likely be misinterpreted as a climb, and be compensated for by pushing the stick forward. This would result in a ‘graveyard spiral’ to planet earth, similar to what JFK Jr likely experienced.

    So while obvious why mammals have this vestibular ‘inertial detection’ system in place, and that its development appears to vary according to need [selective pressures met], what interests me the most regarding its structure is the it has three axiis, each at 90 degrees (referred to at Autonetics as quadrature conformance), as to how this spatial detector ass’y formed initially. I know, a series of random mutations, and convergently evolved in unrelated lineages. Uh, what’s the icon for tongue in cheek?

    Whoa, I hear cheers … ‘curiosity’ has just landed successfully on Mars!

  17. BillyJoe
    Posted August 6, 2012 at 4:46 am | Permalink

    “A city boy goes to the country and sees a bunch of hogs slopping noisily and messily at their trough. “No wonder they call them pigs!”, he exclaims.”

    We have a similar joke. Galahs have a tendency to flock close to or even on roadways. So, when you near miss a group of them, you exclaim “No wonder they’re called galahs.

  18. BillyJoe
    Posted August 6, 2012 at 5:00 am | Permalink

    Today I met Mr and Mrs Darwin and their son.

    I asked them if they were tempted to call their son Charles. This started a long discussion the jist of which was that the mother had done some initial research into the family geneology to see if her husband is, in fact, related to Charles Darwin. She found images of young Charles Darwin which, she says are splitting images of her son and her husband’s father when he was young.

    Unfortunately they are moving to Japan for four years so it will be some time before I get any follow up.

  19. Posted August 6, 2012 at 6:24 am | Permalink

    How can they move so slowly? Don’t they ever need to flee from predators? Do they even have any natural predators? If so, what are their defenses? What strange, interesting animals!

    • MNb
      Posted August 6, 2012 at 9:19 am | Permalink

      Camouflage and claws, mainly camouflage. They are hard to spot in trees exactly because they move so slowly.

  20. W.Benson
    Posted August 6, 2012 at 6:25 am | Permalink

    Of the three reasons (mutation, energy reallocation, reduction of harm) for reduction in vestigial organs, Darwin proposed the two latter mechanisms in 1859. Of course, Darwin knew nothing of how genes and mutations worked. Curiously not long afterwards, Ernst Haeckel (in the History of Creation if I remember correctly) made some very modern statements on the reallocation of energy leading to natural selection for trait loss. This may be surprising to some, since Haeckel has been widely depicted (self-servingly by someone called Steve) as a Lamarckian, whereas in fact he was possibly a stronger proponent of natural selection than Darwin himself.

  21. tualha
    Posted August 6, 2012 at 8:36 am | Permalink

    Obligatory sloth cartoon:


  22. Posted August 6, 2012 at 9:51 am | Permalink

    Yeah sure but, anyways who reads DARWIN anymore, huh?

  23. Launcher
    Posted August 6, 2012 at 12:52 pm | Permalink

    Very interesting posting! I study the neurophysiology of the auditory and vestibular systems (but certainly not in the sloth!), so I thought I’d add my two cents:

    > Well, sloths still have to hang in trees and maintain balance when they defecate … but maybe they don’t need a perfectly-formed inner ear to do that…

    Very slow rotational movements and static position aren’t encoded by the semicircular canals, even in fast-moving animals, because of the mechanical dynamics of the fluid and hair cell receptors. But the otoliths (vestibular organs that are also in the inner ear) as well as visual and proprioceptive feedback would give the sloth the information it needs about body position and its static/slowly moving environment (i.e. with respect to the animal’s head, as one of the most important roles of the SCCs is to control eye movements during head rotations in order to maintain visual fixation). I’d be curious if these other perceptual systems are better developed in the sloth than in its faster moving relatives.

    > What remains is a more definitive test of whether variation in the inner ears of sloths really does matter less than such variation in other species. I’m not sure how one would test that, for you can’t just go in and just manipulate the inner ear.

    There are ways to manipulate the inner ear in the laboratory, such as by block the fluid movement in one or more canals (ENT surgeons do this in human patients). But a simpler experiment would be to study the vestibular-ocular reflex (VOR) in an intact animal. This would amount to measuring eye movements in the dark (i.e. in the absence of visual input) as the animal is rotated. Our working hypothesis would be that the gain of the vestibular-ocular reflex is low and perhaps more variable in adult sloths than in other species. Two-to-one odds that juvenile sloths have a higher VOR gain, as they have smaller heads (that presumably rotate faster) and could make use of the canal’s sensory input.

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