Oh, no, that was a terrible opening. You’ll only know what the heck I’m talking about if you remember JJ from the television show Good Times, and it’s such a pathetic joke it’s only going to appeal to grade schoolers. So if you’re a time-traveling 8 year old from the 1970s, you’ll appreciate the reference. How many of those are reading this right now?
Maybe this will work better. Here’s a small chip of shale I keep at my desk.
My son Alaric and I collected that on a trip to Delta, Utah over 20 years ago. We had permission from the owner of a commercial dig site to rummage around in their tailings*, and we ambled about picking up chunks of rock and splitting them with a hammer. Everywhere we looked were trilobites. We brought home a good haul, chiefly Elrathia, like that one, and lots of Peronopsis. I keep it at my desk as a token of a good memory, and also because it’s about half a billion years old.
I can reach over and touch a half billion year old fossil at will, which I find to be an awesome thrill. That it’s also from a subphylum that was so successful, swarming in our oceans for about 300 million years, yet that ended so finally in the Permian extinction, is humbling. Puny ephemeral humans — we can only dream of achieving the glories of the Trilobite empire.
If you want to learn more about trilobites, I can’t recommend Richard Fortey’s book, Trilobite: Eyewitness to Evolution, highly enough. It’s an excellent, enthusiastic, readable overview of the group. There’s also a gorgeous online guide to the orders of trilobites that’s full of fossil photos and detailed information. But I also recently stumbled across a review paper by Nigel Hughes that looked at them from the perspective of development — O Rhapsody! It’s beautiful!
Despite being extinct for 250 million years, and despite being nothing but fossils, we still have a pretty good idea of the development of trilobites, because they were so numerous and we can find great drifts of entire populations of the animals embedded in lagerstätten. That allows us to see the range of variation and the distribution of different developmental stages, and further, because they’re arthropods, we can see well preserved cuticles of both intact animals and molted shells. And with almost 300 million years of recorded species, we’ve also got a good picture of their evolution. This is a classic evo-devo story.
So, quick, here’s a general introduction to trilobite anatomy. First thing to know is that the ‘three lobes’ of the word ‘trilobite’ refer to the longitudinal divisions of the animal: a central axis with a lateral or pleural lobe on either side of it. There are also, usually, three transverse divisions: cephalic (head) segments, thoracic segments in the middle, and a pygidium or tail.
Not usually shown are the limbs. If you flip over a trilobite, you discover that each segment, except the anterior- and posterior-most, has a pair of biramous appendages — they’re branched legs, with one branch functioning as the walking limb, and the outer branch being lamellate (thin and flat) and probably functioning as a gill. They’re surprisingly uniform and consistent in general structure, from head to thorax to pygidium. One of the curious features of trilobites is that most species are marked by this homonomous condition (that is, maintaining identity or close similarity between adjacent segments), while most of the extant arthropods are strongly heteronomous, making strong distinctions in the structure of adjacent segments.
Now here’s the cool bit: a generalized staging series for trilobites. There are some broad terms for different stages — protaspid, then meraspid, then holaspid — but this diagram makes it clear that growth was by sequential addition of new segments to the posterior end of the animal. This is not an unusual pattern: vertebrates also build segments sequentially from front to back, as do many insects (the short germ band insects), but others, long germ band insects like flies, build the whole collection nearly simultaneously.
Development is the foundation of evolutionary change, and I can’t help but wonder how this pattern, and the unknown genetic constraints behind it, affected trilobite evolution. The early history of arthropods seems to be one of exuberant exploration of the potentials of that modular segmental organization, with trilobites tending to be more conservative than other arthropods. What that means is tricky to interpret: the more inventive arthropods still have descendants around, while trilobites are extinct without issue. But 300 million years is still a fantastically good run, and clearly they had the flexibility to survive major changes in geological history.
The real mystery is why the clade as a whole began to decline after the Ordovician, and how the end of the Permian could so thoroughly quench this gigantic group.
Hughes NC (2007) The Evolution of Trilobite Body Patterning. Annu. Rev. Earth Planet. Sci. 35:401–34.
*By the way, I recommend digging in fossil beds as a great way to connect with the history of the planet with your kids. You can’t make it to Delta? There are quarries that will sell you crates of unprocessed rock, 30 pounds for $75, and you can take them apart in your back yard.