Transitions: The Evolution of Life

The pictures below are all of primates.

The jury is still out as to whether the tree shrew (pictured below) is a primate, or is related to the primates.

Primates come in all shapes and sizes, so the question “how do anthropologists and paleontologists” is a natural one. The question necessarily involves a great deal of comparitive anatomy but I have tried to keep it to a minimum. Readers who need an overview can read this article on the anatomy of the skull. I have written this post in the form of a question and answer session.

What is a primate?
That is actually a good question and the answer is quite complicated.

What do you mean? Aren’t monkeys monkeys?
Well yes, but it’s more complicated than that. It is easier to define modern living primates. Primates as a group do share some unique, universal (among primates) features not shared by other mammals. Unfortunately, the also share features in common with other mammals.

Could you give an example of a unique (or diagnostic) feature that separates one mammal group from another?
Sure! The double pulley configuration of the astralgus (a bone in the hind limb) is diagnostic of artiodactyls.

But, what about primates?
That is a good question. Mivart first defined the order primates. His defination was:

Unguiculate, claviculate placental mammals, with orbits encircled by bone, three kinds of teeth, at least at one time of life; brain always with a posterior lobe and calcerine fissure; the innermost digit of at least one pair of extremities opposable; hallux with a flat nail or none; a well developed caecum; penis pendulous; testes scrotal; always two pectoral mammae.

Wow, that’s a lot!
Actually there is more. Mivart gave his definition in 1873. In 1959 Le Gros Clark added to it:

Preservation of generalised limb structure with primitive pentadactyly (five fingers). Enhancement of free mobility of the digits, especially of the pollex and hallux (both used for grasping). Replacement of sharp, compressed claws by flat nails; development of verysensitive tactile pads on the digits. Progressive shortening of the snout. Elaboration of the visual apparatus, with development of varying degrees of binocular vision. Reduction of the olfactory apparatus. Loss of certain elements of the primitive mammalian dentition. Preservation of a simple molar cusp pattern. Progressive expansion and elaboration of the brain especially of the cerebral cortex. Progressive and increasingly efficient development of gestational processes.

That seems pretty thorough. Is there more?
Yes, there is.

I was afraid of that.
Please don’t interupt. In 1967 Napier added two more:

Prolongation of postnatal life periods. Progressive development of truncal uprightness leading to a facultative bipedalism.

That’s a lot of information, where did you get it?
Mainly from R. D. Martin’s paper “Primates: A Definition”

So what’s the problem with the above definition of primates?
There are two problems. First, some of the above are actually trends, some of which are not features. Instead they refer to developments found only in some members of the group (remember, we are not trying to trace ancestor-descendent relationships at this point. We are trying to provide a definition of an order of mammals). Second, some of these are either traits that are probably primitive features of placental mammals or have arisen by convergence.

So, then how do we define primates?
Martin choose to examine living primates with an eye to creating a new definition, which I won’t bore you with since it is rather long.

You said the definition applies to living primates, what about fossils?
For fossils the definition has to be modified somewhat, but first we have to talk about tree-shrews.

?Tree-shrews?
Yes, you see Le Gros Clark argued that the Tupaiidae are more closely related to primates than to any other placental mammal and should be included in the order primates. This has been argued about ever since. Martin used tree-shrews as a test case for his definition of primates and decided (correctly, I think) that they were not primates.

That’s a pretty scientific approach!
Yes, it is. Paleoanthropology has a well developed scientific methodology and a rich body of theory to draw on.

So what about the fossils?
Since we have only skeletons to examine the definition has to be contracted somewhat. This is what is left:

Well developed, divergent hallux with flat terminal phalanx in the foot. Elongated distal segment of the calcaneus. Relatively large, convergent orbits with restricted interorbital distance. Postorbital bar present; ethmoid exposure in the orbit possible (depending in interorbital distance relative to skull size). Petrosal bulla. Relativly large braincase. Sylvian sulcus on endocast. Dental formula maximally 2.1.3.3/2.1.3.3. Premaxilla short; upper incisors arranged more trnsversly than longitudally. Molars with low, rounded cusps. Lower molars with raised, enlarged talonids.

So, does it identify fossil primates?
Yes, it does. According to this definition omomyids and adapids, for example, are primates.

What about plesiadapids?
The jury is still out on this issue. But see the next post.

For Further Reading:

Primate Adaptation and Evolution”

Primate Evolution: An Introduction to Mans Place in Nature

Primate Evolution

The Evolution of Primate Behavior

Major Topics in Primate and Human Evolution

Additionally, if you have access to a good (University) library:

The American Journal of Physical Anthropology
The Journal of Human Evolution
Folia Primatologica