Tuesday 2 November 2010

What is a Mammal?

I'm going to briefly pause from looking at recent mammalogical news to examine a more general question about mammals. And they don't come much more general than "what is a mammal?"

Doesn't sound a very difficult question, does it? A mammal is a warm-blooded, air-breathing vertebrate that, crucially, feeds its young with milk from its mammary glands. Right? Well, kind of...

For the last twenty years or so, animals have been placed into groups on the basis of something called cladistics. Essentially, the idea is that a proper, meaningful, group of animals will be one that contains a single common ancestor and all of its descendants. Or, to put it another way, that everything in the group has to be more closely related to other animals in that group than it is to anything else. Which seems pretty straightforward and common-sense, and, indeed, is a very useful way of doing things. But applying it strictly does sometimes lead to some fairly surprising results.

Vertebrates first freed themselves entirely from the water when they evolved a way of making their own little pools of water, surrounded by a protective shell, and leaving their tadpoles inside that pool with a supply of yolk to feed off until they became developed enough to hatch. The animals that evolved this feature are called amniotes. Later on, some mammals evolved a way of doing away with the shell, and keeping the pond inside a membranous sac in the mother's body, but the principle is the same, and they still count as amniotes.

Not long after the amniotes first appeared, they split into two great evolutionary lines. One led to the reptiles and birds, and the other, called the Synapsida, led to the mammals.

Wait a minute, I can hear you saying, but didn't mammals evolve from reptiles? Well, it depends what you mean by "reptile". Certainly, if we could look at the early synapsids today, most people would probably call them reptiles. They were cold-blooded, hairless, laid eggs, wouldn't have produced milk, and anyway, they just kind of looked reptilian. Take a look at this one, for example. By most people's standards, that's a reptile.

But here's how the early amniotes evolved into the animals we have today:

Crocodiles    Birds      Other      Turtles      Mammals
    ^           ^          ^           ^            ^
    |           |          |           |            |
    |           |          |           |            |
    -------------          |           |            |
          |                |           |            |
          |                |           |            |
          ------------------           |            |
                   |                   |            |
                DIAPSIDA            ANAPSIDA    SYNAPSIDA
                   |                   |            |
                   ---------------------            |
                             |                      |
                            (A)                     |
                             |                      |

Bearing in mind out definition for what constitutes a "group" of animals (a monophyletic clade if we want to get technical - but let's not) there are two problems with this chart. The first is that reptiles aren't actually a proper group at all - they can't be, because crocodiles are more closely related to birds than they are to, say, turtles. Which isn't to say that crocodiles are particularly close to birds, admittedly, just that they're even further from turtles. (This is because birds evolved from dinosaurs, which were fairly close to crocodiles).

When we say that a proper biological group has to consist of a common ancestor, and all of its descendants, the key word is "all". The common ancestor of all living reptiles is at the point I've marked (A), and if we want to include all of its descendants, then we have to count the birds. Either birds are reptiles, or reptiles don't really exist as a meaningful group. Bummer!

Be that as it may, the other problem is that even if the reptiles are a group, their last common ancestor, and therefore the first reptile, is the creature at (A), and mammals didn't evolve from that. So mammals evolved from creatures that certainly looked very reptilian, but which weren't, in a strict scientific sense, reptiles as we understand them today. They are entirely their own line.

But, at any rate, it's fairly clear that while the early synapsids may not have been reptiles, they weren't mammals, either. At some point, they evolved into mammals, and all of the earlier forms of reptilian-looking synapsids died out. So at what point did that happen? When did not-mammals become mammals?

The obvious answer is "when they developed mammary glands and began producing milk". Which is all very well, but a bit of a bugger when all you've got to go on is fossil bones. How do you tell from the bones whether the animal produced milk or not? You can't, pretty much. So palaeontologists have to use a different definition.

Skull of a turtle - note that the lower jaw consists of at least four different bones. (d = dentary, ar = articular)
The lower jaws of the early amniotes consisted of multiple bones. For example, there was the dentary bone, which had most of the teeth, and the articular bone, which formed the hinge joint with the skull. In the line that led to reptiles and birds, this didn't change much (at least until birds evolved beaks), but in the synapsids, something strange began to happen. The dentary bone began to get larger, slowly pushing out and shrinking the other bones, until most of them disappeared altogether.

Eventually, the dentary formed its own joint with the skull, and the lower jaw actually had two joints on each side for a while. That's not much of a problem, so long as the joints are lined up properly, but there's really no need for it, so eventually, the only remaining other bone in the lower jaw, the articular, began to shrink as well. In the end, the dentary was the only bone left in the lower jaw at all; and in mammals, we call it the mandible.

Lower jaw of a mammal - note the absence of separate bones
But the articular bone, and its old joint with the skull didn't disappear altogether. Both it, and the skull bone that it used to attach to shrank and became entirely separated from the jaw, moving up the side of the head. They are still there today, still with the old joint between them, but now we call them the malleus and incus, and they form two out of the three bones in the middle ear.

And that, at least when you're looking at fossils, is the defining characteristic of a mammal: that it has one bone on  each side of the lower jaw, and three bones in each middle ear.

[Pictures from Wikimedia Commons]


  1. Are amniotes limited to the mammalian evolutionary path, or would they include viviparous snakes and sharks?

  2. Amniotes are the group that includes reptiles, birds, and mammals - everything that's descended from that animal that first evolved an egg that could survive on dry land. Some of them, such as most mammals, and viviparous reptiles (such as sea snakes), no longer lay eggs, but keep the embryo inside the mother's body. In fact, the word 'amniote' refers to the 'amnion', a special membrane that encloses the embryo in all of these animals, whether or not there is a solid shell outside of it. Its part of the afterbirth in mammals, but its also that thin lining you get inside of (for example) a chicken egg.

    So all reptiles, birds, and mammals are amniotes. Viviparous sharks and amphibians don't count, because they have evolved different and evolutionarily separate ways of doing what they do, and represent earlier branches from the great vertebrate tree.