OH, GIVE ME A HOME, WHEN THE TRILOBITES ROAMED
Trilobites tell me of ancient marine shores teeming with budding life, when silence was only broken by the wind, the breaking of the waves, or by the thunder of storms and volcanoes. The struggle of survival already had its toll in the seas, but only natural laws and events determined the fate of evolving life forms. No footprints were to be found on those shores, as life had not yet conquered land. Genocide had not been invented as yet, and the threat to life on Earth resided only with the comets and asteroids. All fossils are, in a way, time capsules that can transport our imagination to unseen shores, lost in the sea of eons that preceded us. The time of trilobites is unimaginably far away, and yet, with relatively little effort, we can dig out these messengers of our past and hold them in our hand. And if we can learn the language, we can read the message.
RICCARDO LEVI-SETTI, TRILOBITES
AMBASSADORS OF DEEP TIME
One of the most popular of all fossils for amateur collectors and professional paleontologists alike are the trilobites. These creatures lasted from 550 to 250 million years ago and, over those 300 million years, evolved more than 5000 genera and 15,000 species, all of which are now extinct (figure 4.1). They range from the tiny Acanthopleurella (barely 1 millimeter [0.04 inch] in length) to the giant Isotelus rex (more than 70 centimeters [2.3 feet] long). Since they are relatively easy to collect in many places, and extraordinarily abundant almost everywhere in beds of the Early Paleozoic (especially the Cambrian), they often become the core of many amateur fossil collections. Their wonderfully complex shapes, elaborate ornamentation, bizarre structures of the eyes and many other parts of their anatomy, and surprising features make them irresistible to most fossil collectors.
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Figure 4.1
Reconstruction of two trilobites as they may have appeared in life. (Courtesy Nobumichi Tamura)
This fascination is not confined to modern times. A trilobite from the Silurian carved into an amulet was found in a rock shelter more than 15,000 years old. A trilobite from the Cambrian preserved in chert was carried a long way by Australian Aborigines and carved into an implement. The Ute peoples used to carve the common trilobite Elrathia kingi, from the House Range of Utah, into amulets. They called them timpe khanitza pachavee (little water bug in stone house). Elrathia kingi are so abundant in this locality that they are commercially mined with backhoes and are sold in huge quantities to nearly every rock shop and fossil dealer around the world.
Even more important, trilobites were the first large shelled organisms on Earth. There is abundant evidence from the genetic divergence times of their close relatives that soft-shelled trilobites were around in the earliest Cambrian and developed mineralized shells in the Atdabanian, the third stage of the Early Cambrian (see figure 3.4). This may be because atmospheric oxygen levels were finally high enough that trilobites could crystallize calcite in their shells. Most of the creatures that preceded them either were soft-bodied, with no hard parts or shells, or had tiny and inconspicuous shells (chapters 2 and 3); therefore, they were fossilized only in environments with conditions that favored preservation, not decomposition (chapter 5). Not only did trilobites have a large complex shell made of the protein chitin (as do crabs, lobsters, shrimps, insects, spiders, scorpions, and all other arthropods), but this relatively soft and easily decayed shell was fortified by layers of the mineral calcite. Thus trilobites were much more likely to be fossilized than any other Cambrian creature, since they were one of the few groups with mineralized shells. The appearance of hard-shelled trilobites in the Atdabanian makes them overrepresented in the fossil record, and they give the false impression that there was a “Cambrian explosion” of life between the Tommotian and the Atdabanian (see figure 3.4). Instead, it was an “explosion” of animals with mineralized skeletons.
The abundance of easily fossilized trilobites in deposits of Late Cambrian age meant that more than 300 genera in 65 families were recognized, completely overwhelming all other fossil groups known from that time. In just about any Cambrian deposit, the majority of fossils are trilobites, so paleontologists use the stages of trilobite evolution to tell time in the Cambrian.
WHAT IS A TRILOBITE?
Trilobites are the earliest known fossilized arthropods, the phylum that includes insects, spiders, scorpions, crustaceans, and many other creatures (chapter 5), and they clearly display all the features of that phylum. Like all other arthropods, they had a jointed exoskeleton that fell apart when they molted, so the fossils are often incomplete pieces of molted shell, and not the complete animal, which likely lived on to molt again. Unlike that of most other arthropods, however, the chitinous exoskeleton of trilobites was reinforced with mineralized calcite, which made them much more fossilizable than insects or spiders or scorpions or most crustaceans.
The “head” of most arthropods is called the cephalon (Greek for “head) in trilobites (figure 4.2A). It is usually a broad structure with two “cheek regions” on each side of a central lobe (“nose”) called the glabella. On each side of the glabella are typically two eyes. Some trilobites had tiny eyes or none at all, so they had limited vision or were blind; others had huge eyes that wrapped around and gave a 360-degree range of view to spot any predators. Many advanced trilobites had lenses made of two crystals of calcite, forming a doublet lens structure that corrected for spherical aberration in thick lenses. About 400 million years after trilobites evolved these features, they were reinvented by Christian Huygens, the great Dutch scientist, in the sixteenth century. Even more important, trilobites were probably the first creatures on Earth to have true eyes and to use visual clues to find food and avoid predators.
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Figure 4.2
Basic anatomy of a trilobite: (A) top view of the complete exoskeleton; (B) bottom view of the cephalon; (C) cross section through the axis, with skeletal parts indicated in black. (Modified from several sources)
The cheek regions broke off from the central part of the cephalon (cranidium) during molting, so most trilobite fossils consist of just the center of the cephalon. A good trilobite specialist often needs only a cranidium to identify the species. There are great variations in the details of the eyes, the glabella, the shape of the cheeks, and the spines on the edge of the cephalon. On the front of well-preserved specimens are two antennae that trilobites used for feeling their way around in dark muddy waters. On the bottom is a plate that partially covered the mouth, with mouthparts that were used for sucking up food-rich mud and digesting the nutrients out of it (see figure 4.2B). Most trilobites were deposit feeders or mud grubbers.
The middle part of the body of a trilobite is called the thorax, as in most arthropods. In trilobites, the thorax is divided into segments, allowing the middle of the body to flex and curl as the animal moved or to roll up for protection. Each segment has two lobes on the sides (pleural lobes) and one that runs down the central axis (axial lobe). It is this side-by-side division of the body into three lobes that gives the group their name. Some trilobites have just two or three thoracic segments, so they were not very flexible and must have lain flat nearly all the time. Others have many segments, so they could roll up tightly into a ball to deter predators, such as the isopod crustaceans known as “roly-polies” or “sowbugs” do today. Well-preserved fossils show that beneath each thoracic segment was a pair of walking legs, and on each side a pair of feather-like gills attached to the base of the legs.
The tail end of the trilobite is not called the abdomen, as in many arthropods. Instead, it is known as the pygidium (Greek for “little tail”). In most trilobites, the last few thoracic segments are fused into a single large plate-like pygidium. The olenellids, however, are very different.
OLENELLUS AND THE FIRST TRILOBITES
Once you acquire a discerning eye for telling trilobites apart, Olenellus is one of the easiest trilobites to recognize (figure 4.3). It was named and studied in detail by none other than the pioneering Cambrian expert Charles Doolittle Walcott, in 1910 (chapter 1). Its most distinctive feature is the lack of fusion of the last few segments of the thorax into a single pygidium. Instead, Olenellus has a long spike on the tail. This is a very primitive feature, which is no surprise, since the olenellids are the oldest trilobites known.
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Figure 4.3
A specimen of Olenellus, showing the characteristic D-shaped cephalon, bulbous tip of the glabella, large crescent-shaped compound eyes, spines on the tips of the cephalon and on certain thoracic segments, and absence of a large fused pygidium, or tail segment. (Photograph courtesy Wikimedia Commons)
In addition to the absence of a pygidium, there are many other distinctive or primitive features in Olenellus. The cephalon is large and shaped roughly like a capital D. There is no line of rupture (cranial suture) on the top of the cephalon, which separated the cheeks from the cranidium when Olenellus molted. Two big crescent-shaped eyes wrap around each side of a furrowed glabella, which has a bulbous knob at the tip in front. The eyes are simple, with many tiny lenses made of calcite rods packed closely together, like the typical compound eyes of most insects and many other arthropods. The eyes could not have formed a clear photographic image, but would have alerted the trilobites to large areas of light and darkness and movement near them. Studies of the extraordinary Cambrian faunas such as those preserved in the Burgess Shale in Canada and the Maotianshan Shale in China show that there were almost no large predators at that time (chapter 5). The largest may have been the 1-meter (3.3-foot) long Anomalocaris. Fossils show that it clearly took bites out of the trilobites found in the Middle Cambrian Burgess Shale, where it was discovered. But compared with the later Paleozoic, there was not a lot of predation pressure, and trilobites were relatively simple and unspecialized in the Cambrian. Not until the Ordovician do we get super-predators, such as nautiloids with shells over 6 meters (20 feet) long. Only then did trilobites evolve distinctive shells specialized for burrowing or swimming or rolling up into to a ball as a defense against tougher predators.
One other striking feature of Olenellus is that it is very spiny on the edge of its shell. There is typically a spine (genal spine) sticking out of the back corners of the cephalon. Many have broad spines sticking out of their thoracic segments, and then backward, usually segment number 3. Some have additional spines protruding from the front of the cephalon as well. These spines often help paleontologists recognize different genera and species within the olenelloids.
Once olenelloids appeared in the Atdabanian stage of the Early Cambrian (about 520 million years ago), they flourished into multiple genera and species found almost everywhere around the world in that stage and in the succeeding Botomian stage of the Early Cambrian. They then vanished at the end of the Toyonian stage (about 509 million years ago). Bruce Lieberman of the University of Kansas has analyzed thousands of specimens of olenellids and concluded that their ancestors originated in what is now western Russia or Siberia at the beginning of the Cambrian, but, like other trilobites, were not calcified or fossilized until the Atdabanian stage.
WHAT HAPPENED TO THE TRILOBITES?
Through the rest of the Paleozoic, the trilobites were hammered by a series of mass extinction events (figure 4.4). They include the multiple minor extinctions in the Late Cambrian, when several pulses of disasters wiped out the diversity of trilobites, wave after wave. During the Ordovician, trilobites experienced their first encounter with large predators, probably gigantic nautiloids. Trilobites quickly became more specialized and easier to tell apart as they soon adopted a variety of shapes and lifestyles that made them less vulnerable to predation. These adaptations included burrowing (the smooth “snowplow” trilobites known as Asaphida or Illaenida), rolling up into a ball (the Calymenida), or becoming tiny (the Trinucleida, such as Cryptolithus, the thumbnail-size “lace collar” trilobite). Then came the Late Ordovician extinction (about 450 million years ago), and only a few lineages survived into the Silurian and Devonian. The final flourishing of trilobites occurred in the Devonian, when the complex-eyed Phacopida were common, along with large spiky trilobites like Terataspis (about 0.5 meter [1.5 feet] long). The Late Devonian extinctions at 375 million years ago and 357 million years ago wiped out all but one order of trilobites, the relatively small and simple Proetida, which persisted in the background for another 125 million years.
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Figure 4.4
Diversification and extinction of the trilobites. (From several sources)
Finally, the trilobites disappeared during the great Permian extinction, some 250 million years ago, the largest mass extinction in Earth’s history, when 95 percent of all marine species vanished. This huge event wiped out not only the last of the stragglers among trilobites, but also the two dominant groups of Paleozoic corals (the tabulates and the rugosids), as well as the blastoids (relatives of the crinoids, or “sea lilies”) and the fusulinid foraminiferans (incredibly abundant protozoans with shells shaped like rice grains). There have been many controversies about what caused “the mother of all mass extinctions,” but current data indicate that the largest volcanic eruption in geological history, the Siberian lava flows, helped trigger an extremely rapid greenhouse climate that made the oceans too hot and acidic to support much life, and overcharged the atmosphere with too much carbon dioxide and not enough oxygen. These, along with some other catastrophic events, destroyed all but a tiny percentage of life on Earth.
SEE IT FOR YOURSELF!
A number of museums have trilobite fossils on display, although most do not show very good complete specimens of olenellids. Some that do include the Denver Museum of Nature and Science; Field Museum of Natural History, Chicago; Geology Museum, University of Wisconsin, Madison; and National Museum of Natural History, Smithsonian Institution, Washington, D.C.
  EMIGRANT PASS, NOPAH RANGE, CALIFORNIA. Take Interstate 15 to Baker, California; leave the interstate; and drive north for 48 miles on California State Route 127 (Death Valley Road) to Old Spanish Trail. Turn right on Old Spanish Trail and proceed through Tecopa to Emigrant Pass. The exposure is just to the west of the summit of the pass on the south side of the road (GPS coordinates = 35.8856N, –116.0603W). Two good Web sites are “Trilobites in the Nopah Range, Inyo County, California” (http://inyo.coffeecup.com/site/cf/carfieldtrip.html) and “Ollenelid Trilobites at Emigrant Pass, Nopah Range, CA” (http://donaldkenney.x10.mx/SITES/CANOPAH/CANOPAH.HTM).
  OAK SPRING SUMMIT, LINCOLN COUNTY, NEVADA. From Caliente, Nevada, take U.S. Route 93 west for 10 miles, or east for 33 miles from the junction with Nevada State Route 375 and 318 (between Hiko and Ash Springs). Look for a turnoff on the north side of the highway, signaled by a prominent Bureau of Land Management sign that reads “Oak Springs Trilobite Site.” Turn north, drive along the dirt road, park in the gravel lot, and then hike up the Trilobite Trail from the trailhead across the sagebrush until you arrive at a flat area covered with pieces of the Pioche Shale. Turn over shale, and you will find many fine cephala, occasionally better specimens, of every age and size. A good Web site is “Oak Spring Summit” (http://tyra-rex.com/collecting/oaksprings.html).
FOR FURTHER READING
Erwin, Douglas H., and James W. Valentine. The Cambrian Explosion: The Construction of Animal Biodiversity. Greenwood Village, Colo.: Roberts, 2013.
Fortey, Richard. Trilobite: Eyewitness to Evolution. New York: Vintage, 2001.
Foster, John H. Cambrian Ocean World: Ancient Sea Life of North America. Bloomington: Indiana University Press, 2014.
Lawrance, Pete, and Sinclair Stammers. Trilobites of the World: An Atlas of 1000 Photographs. New York: Siri Scientific Press, 2014.
Levi-Setti, Ricardo. The Trilobite Book: A Visual Journey. Chicago: University of Chicago Press, 2014.