Sunday 28 November 2010

Telling fossil species apart - a brontothere's nose

Brontotherium skull - or should that be Brontotherium prousti?
When we think of specific kinds of living animals, we tend to think of individual species, at least where those animals are fairly familiar to us. But prehistoric animals are almost always referred to just by their genus name. A proper species name has two parts to it (such as Homo sapiens), but the genus name is just the first part of that, and that's how we think of most prehistoric creatures. Consider the dinosaurs, such as Triceratops, Velociraptor, and Diplodocus, or mammals such as Smilodon, the sabre-toothed cat. Probably the only exception - and arguably one of only two creatures whose common name is the same as its scientific name - is Tyrannosaurus rex. (The other one is Boa constrictor, if you're wondering).

A genus is a group of closely related species. There are many genera that only contain one species, but most have a number of different species at any given point in time, and even more when you look over the whole of their evolutionary history. It's as if we thought of the living genus Canis as a single entity, ignoring the fact that it includes wolves, coyotes, and jackals, never mind extinct species such as the dire wolf.

There's a reason for this, of course, which is that it's really tricky to tell actual species apart from their fossils. Taking the traditional definition of a species as something that doesn't cross-breed with other species to produce fertile hybrids (and the modern definition is a bit more complicated than that), how on Earth would you know, if you're only looking at a fossil? Instead, the definition more commonly used for fossil animals is that a species is a group of specimens so similar that there's no way to consistently tell them apart, even when you have a complete skeleton. But most fossils aren't complete, or are at least damaged by several million years of being squished under tons of rock, which makes it very difficult to spot the tell-tale differences between the equivalents of coyotes and jackals.

Sunday 14 November 2010

The Polar Bear's Bite

Bears are generally omnivorous animals, and will eat pretty much whatever is available. Although they look fairly fearsome, and do, of course, eat meat, they also consume a relatively large amount of plant material, such as berries, leaves, grass, nuts, and so on. This is useful, especially for a large animal, since they are unlikely to go very hungry for long, and plants are always more readily available than animals.

The polar bear (Ursus maritimus) is, of course, an exception. Unlike all other living bears, they feed exclusively on meat, and really don't eat plants at all. They are, to use the technical term, hypercarnivores, more similar in this respect to cats than they are to their own relatives. Nor do the differences end there. Most obviously, they are adapted to bitterly cold environments, and, indeed, they soon get heat stroke if taken to places that most other bears would be perfectly happy - anything much above 10°C (50°F) makes them uncomfortable. Not only that, but they are also much better swimmers than other bears, and have claws that grip into the ice when they walk.

Given these differences, you might think that polar bears represent an early branch from the lineage that led to all the other bears, one that adapted itself to both an extreme environment and a (for bears) unusual diet. But that turns out not to be the case, with polar bears having diverged from their closest living relatives, the brown bears, long after that line split from the other species.

 Polar      Brown     Sun Bear     Black
 Bear       Bear                   Bears
   |          |          |           ^
   |          |          |           |         Sloth Bear
   ------------          -------------             |
        |                      |                   |
        |                      |                   |
        ------------------------                   |
                   |                               |
                   |                               |

Rather more surprising, perhaps, is just how recently this happened. Genetic analyses show that the first polar bears evolved no more than 0.7 million years ago, which really isn't very long in evolutionary terms - in contrast, most other species of bear diverged around 5 million years ago. In fact, its quite probably a lot less than 0.7 million years. The oldest known skeleton of a polar bear is only around 130,000 years old, which is sufficiently young to retain a good deal of DNA.When that DNA was analysed earlier this year, it seemed to show that the animal in question was so incredibly close to brown bears that it probably lived at around the time that the two diverged, and would therefore have been one of the first members of its species - a rare find, indeed.

What this means is that, after they first evolved, polar bears must have undergone a very rapid sequence of changes, quickly adapting themselves to their environment and new diet. Judging from the shape of their skulls, they must have changed at least twice as quickly as any other bear species did, and that's probably an underestimate.

How has the polar bear managed to adapt to a new diet in such a short time? Generally speaking, hypercarnivores such as lions tend to have strong skulls with powerful jaw muscles and large shearing teeth, suitable for slicing through tough meat and bone. This is much less true of bears, adapted as they are (in most cases) to a more omnivorous diet. But, compared with brown bears, polar bears have a long, rather sleek skull, which is even weaker than the heavy rounded skulls of their relatives. This is probably, at least in part, to make it easier for them to nuzzle through holes in the ice to catch seals.

Some recent stress analyses using the sort of computer models normally used in engineering design have shown that while the jaw muscles of polar bears are just as strong as those of normal bears, the skull itself is less able to take up the strain of a powerful bite. Granted, that's assuming the computer models are right, but even so, it does seem a bit odd. Similarly, while they do have less premolar teeth than many other bears (a feature they share with cats, for example), the teeth overall seem somewhat weaker than one would expect for a purely carnivorous animal.

One reason may be what the polar bears are eating. Their diet consists almost entirely of seals, which, unlike the antelopes eaten by lions, have a lot of soft blubber. If their food is less tough than the meat favoured by other hypercarnivores, they don't need such strong jaws, and the evolutionary change required to adapt to that diet may be less than it first appears. It may also help that, while animals like leopards and wolves regularly take down prey  larger than themselves, polar bears obviously don't. Seals just don't get that big!

There is, however, a clear downside to this rapid evolution and specialisation. As ice sheets melt, the habitat of polar bears is shrinking. But what's bad for them can be quite good for their relatives, the brown bears. Brown bears (Ursus arctos) are a diverse species, including both the grizzlies and Kodiak bears of North America, and the "common" brown bears of Europe and Russia, among others. They're pretty adaptable, and can eat a wide range of food, making them at home in dense forests, open woodland, or even tundra. As the polar bears' preferred environment shrinks, brown bears can move steadily further north.

The sort of things that brown bears eat are often tougher than seal blubber, and their teeth and bites are accordingly stronger. This means that, even if polar bears were to start eating such things (which isn't, perhaps, very likely in the first place), the brown bears would still do better. Brown and polar bears are still so closely related that they can interbreed to produce hybrids, at least some of which are apparently fertile. If that continues to happen, and the brown bears muscle in on the polar bear's territory, out-competing them for food, then it's the browns that will eventually win out, diluting the purebred polars into non-existence.

As if the great white bears of the north didn't have enough problems to be going on with...

[Picture from Wikimedia Commons]

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]