A common argument leveled against the theory of evolution is that scientists have not been able to produce the expected transitional fossils that show the change of one species into another. If evolution were true, wouldn’t there be instances of clear intermediary species, like, for example, a species that was half whale and half hippo to show the transition between those two? In this BioLogos podcast, Kelsey Luoma addresses this misconception about what a transitional fossil actually is. Rather than a mix between two related species, transitional fossils point back to the common ancestors that modern species share. The fact is that the number of transitional species is massive and it grows with each passing year. Given the rarity with which organisms are actually fossilized, the amazing thing is actually the completeness of the fossil record, not its incompleteness. The transitional species story strongly supports, and certainly does not disprove, evolutionary theory. ¹

1. To hear the full audio clips which have been referenced go to:

• Rational Response Debate with Kirk Cameron (from Way of the Masters)
• Behind the Scenes with Dr. Neil Shubin (from Cincinnati Museum Center)
• Mark Norell Publishes New Archaeopteryx Findings (from American Museum of Natural Sciences)
• Texas A&M Professor Discusses Findings of Autralopithecus Sediba and its Relationship to Humans (from Texas A&M University)
• Intro/outro music composed by Martin Minor (Minor2Go).

An audio only version of the podcast can be downloaded here.

Transitional Fossils

Some time ago, the Discovery Institute’s Casey Luskin commented on the human origins exhibit at the Smithsonian Institution, suggesting that palaeoanthropologists use evolutionary theory to describe the progression of the human lineage even when they don’t have transitional fossils with which to work. He writes:

What's ironic, however, is that if you ask the question How Do We Know Humans Evolved? the answer you’re given is, “Fossils like the ones shown in our Human Fossils Gallery provide evidence that modern humans evolved from earlier humans.” So whether you find fossils or you don’t, that’s evidence for evolution.

Indeed, it has become an article of faith for those espousing both the young earth creation (hereafter YEC) model and many who hold to the intelligent design model that transitional fossils do not exist and therefore evolution has not taken place. Support for this position usually entails attacking the weak areas of the fossil record, where burial processes have left us little with which to work, or the creation of straw men arguments in which transitional fossils are defined in such a way that none could ever be found. Often this centers on the concept of “missing link,” a term that is habitually used in the popular press and young earth creation and intelligent design literature when referring to fossil remains but which has little to no meaning for biologists or palaeontologists. As Ahlberg and Clack (Ahlberg and Clack 2006) write:

But the concept has become freighted with unfounded notions of evolutionary ‘progress’ and with a mistaken emphasis on the single intermediate fossil as the key to understanding evolutionary transitions. Much of the importance of transitional fossils actually lies in how they resemble and differ from their nearest neighbours in the phylogenetic tree, and in the picture of change that emerges from this pattern.

Contrary to common misconceptions, the fossil record does not record one single lineage for any family of organisms but rather a series of branches, with many related species coexisting synchronously. Darwin hypothesized that the evolutionary record reflected this bushiness and drew such a diagram in his journal. At the time, though, he had little in the way of fossil evidence to back up this position. Much has changed since his day.

An analogy for understanding this “bushiness” was best described by Prothero and Buell (Prothero and Buell 2007). They suggest that the reader consider his or her own genealogy. You and your siblings are the direct descendents of your parents and, while you are similar to them, each of you has different characteristics not shared with them as well as characteristics that you do share. Your parents have siblings as well (your aunts and uncles), and your grandparents are their last common ancestors. These siblings have their own children (your cousins), who have different and similar traits relative to their parents. They are broadly recognizable as being related to you (“oh, I see you have Aunt Edna’s nose”) but three or four generations out, they will become less and less so. These are the “nearest neighbours” that Ahlberg and Clack describe. In this analogy, each of these cousins represents a transitional form from what was (your grandparents) to what will be down the road.

For example, no one would confuse a frog with a salamander but if you trace the fossil record of each back in time, eventually you encounter a fossil, Gerobatrachus hottoni which was recently discovered (Anderson et al. 2008) that is best described as a “frogamander,” having the basal characteristics of both frogs and salamanders. Had we seen such an animal at the time, it is likely we would not have found it remarkable because it would have resembled the species around it. One lineage eventually diverged into frogs, salamanders and other amphibians. Most (just like Darwin proposed in his tree diagram with the little hatch marks at the tip of many branches) went extinct.

Taxonomy and the Beginnings of Human Origins

All life is classified based on a system devised by Carolus Linneaus in 1735 in his remarkable work Systema Naturae. This system gives all recognized species an individual place based on a system of hierarchy. The study of classification is known as taxonomy. A taxonomic ranking for humans would be this:

When a fossil is excavated, the first thing that the palaeontologist does is make a taxonomic assessment of where it fits in a sequence of known fossils. Traits that are shared with other like species or genera are referred to as primitive traits. Examples of this in humans are five fingers and the presence of three arm bones. We share this with all mammals. Traits that are new or are not shared with other like species are referred to as derived traits. Examples of this in humans are the skeletal changes in the pelvis and the foot to allow for walking upright. We do not share these with any other primates.

Transitional fossils in the human fossil record are distinguished at both the genus and species level. This group includes the extinct genera Ardipithecus and Australopithecus and the current genus Homo. All species except Homo sapiens are extinct. Much of the recent study of early humans focuses on the transition from Ardipithecus (‘Ardi’) to Australopithecus (‘Lucy’ and similar fossils) and from Australopithecus to Homo, the genus that led eventually to us. While each of the australopithecine species identified in the fossil record has derived characteristics that separate them from their ancestors and from each other, only one led to the genus Homo.

In future posts, I will describe the evidence for human evolution and why this evidence is compelling. It suggests that we have had a long, varied history filled with great leaps of change, crushing defeat, and eventual expansion into all areas of the globe.

Notes

Ahlberg, P. & J. Clack (2006) A firm step from water to land. Nature, 440.

Anderson, J. S., R. R. Reisz, D. Scott, N. B. Frobisch & S. S. Sumida (2008) A stem batrachian from the Early Permian of Texas and the origin of frogs and salamanders. Nature, 453, 515-518.

Prothero, D. & C. Buell. 2007. Evolution: What the fossils say and why it matters. Columbia Univ Pr.

Introduction

One of the reasons that some of us are hesitant to accept evolutionary creation is that it seems to make God responsible for the suffering and death of innumerable creatures over millions of years—before humans ever existed or sinned against their creator. Since we believe in and worship a God who is loving, benevolent, and all-powerful, it sounds quite implausible that our God would have created a world like that; therefore, any scientific evidence for evolution must be incorrect.

Other people look at the scientific evidence for evolution and find a compelling case that it has taken place during our earth's history. On this basis they may conclude that if evolution is true, then the belief in an all-powerful, perfectly good God must be false!

The trouble with both of these views is that they tend to invoke a completely abstract, philosophical god, not the living God of the Bible—the God who became a human being, experienced unimaginable suffering, and died in a grotesque and humiliating public display. The death of Jesus completely defied the expectations (and common sense) of his followers, as well as the expectations of any “rational” understanding of the way the Creator of the universe should act in the world. On the cross, in the person of Jesus, God took upon himself far more suffering than any creature has ever experienced.

If God himself is willing to die, particularly in such a gruesome way, then perhaps we should at least consider the possibility of God allowing the death of other creatures, too. But would this really be compatible with what we know of God through Scripture? In this essay, we will explore this quandary through a “theology of the cross”, a concept articulated by pastor George Murphy in his book Cosmos in the Light of the Cross.¹

Theology of the cross

Before we jump into the theological problems associated with evolution, let’s take a look at how we understand Christian theology itself. For the reformer Martin Luther, any theology (or science) that tries to reach knowledge of God apart from the cross is bad theology.² Instead, Luther pointed to a theologia crucis, in which the true God is seen first and foremost “through suffering and the cross”. To make his point even clearer, Luther insisted that “the CROSS alone is our theology”.³ It is the lens through which we view everything.

Of course Martin Luther, having lived in the 16th century, was not aware of the vast history of life on our planet (or any other aspect of modern science, for that matter), but George Murphy draws from Luther’s teachings the foundation that all human knowledge begins with the Word made flesh and crucified.⁴ With the cross of Christ as the ultimate framework through which we view reality, we are bound to view the processes of nature quite differently. As Murphy explains it,

A theology of the cross is an explication of belief in a God who becomes a participant in the history of the universe and thereby shares in the suffering, loss, and death that are part of worldly experience.⁵

God does not sit idly by and watch unaffected as his creatures suffer, but neither does he swoop in and make everything completely effortless and easy. Instead he chose another way, the crucifixion of Jesus—certainly not the approach that we would have preferred! The apostle Peter went so far as to try to talk Jesus out of it, but he was met with a stern rebuke (Matthew 16:21-23).

Why is evolution so disconcerting to Christians?

The problem of suffering throughout all of human history is troubling enough for us to reconcile with a loving, personal God. But in addition to that, the discovery of vast numbers of fossils reveals that death has taken place on a far greater scale than we had ever imagined. Both the wide variety of extinct creatures and their sheer numbers is quite staggering, and it raises questions about our Creator:

The picture of a God who is immune from suffering and death but who forces organisms through millions of generations and extinction is disturbing to those who believe in a God of love.⁷

The mass extinction of life on earth was already well established by the early 19th century—decades before Darwin’s research—and extinction can be empirically verified independent of any theory of evolution.⁸ The fact that the earth’s crust is a veritable graveyard of long-lost creatures is deeply troubling, and as late as the 1790’s, distinguished intellectuals such as Thomas Jefferson denied the very possibility of extinction.⁹

But in addition to the reality of species extinction, the theory of evolution by natural selection proposes that new species also arise in an environment containing widespread pain and death. Both the creatures that are now living and those that are gone are tainted by an “acrid smell of death”.¹⁰ It makes us wonder, if our Creator is not the God of the dead, but of the living (Mk. 12:27), where is God’s presence in the evolutionary picture?

In all honesty, creation through evolution is not what we would expect from God, but Scripture is full of examples in which God acts in unexpected ways. After all, God’s choosing to undergo an agonizing death on a cross is not what we would expect from the all-powerful Creator of the universe, either. In both cases, new life comes about through pain, suffering, and death. As George Murphy puts it,

A priori ideas about God have to be overcome, and God's character has to be learned from God's self-revelation.¹¹

God’s fullest self-expression is in Jesus Christ himself, one who is intimately familiar with and personally endured creaturely pain and death. The theology of the cross reveals that God's self-revelation takes place in situations of suffering, loss, and apparent hopelessness, much like situations that occur through natural selection.¹²

The crucifixion is disconcerting too

Not only is creation through evolution an unexpected and unsettling process, but so is the crucifixion of Jesus! Killing someone by hanging them on a cross is an unbearably painful, prolonged, humiliating form of death. It was such a horrific type of public execution that it wasn't until after the Roman Empire stopped the practice of crucifixion—and people no longer witnessed it personally—did the cross become a visual object of devotion.¹³ Our culture is sufficiently removed from crucifixion that we are desensitized to its original significance, but to connect it to our current context, imagine the reaction you would get by wearing jewelry designed to look like an electric chair.¹⁴

Once we are more attuned to the brutality of crucifixion, it seems all the more striking that the cross is the sign of God’s work, what George Murphy calls “the trademark of God”.¹⁵ The suffering and death of Jesus is featured prominently in the Gospels, but the crucifixion-resurrection pattern is strongly resonant within the Old Testament, too. Israel suffered and toiled as slaves in Egypt for centuries before they were rescued in the Exodus, bringing life to a people who were spiritually dead. Centuries later, the nation of Israel would experience death again when the Babylonians destroyed the Davidic monarchy, burned their Temple, killed their people, and sent many into exile.¹⁶ Neither Israel (God’s chosen people) nor Jesus (God’s own son) were spared from death and suffering; rather, suffering seems to have been the way in which God re-forms and renews humanity to fully bear His own image.

Redemption extends to all of creation

Fortunately, God’s story does not end with death. God gives new life after his creatures have been subjected to terrible circumstances. Redemption was promised to Israel itself—Ezekiel’s vision of the valley of dry bones describes how God would renew His chosen people (Ezek 37:1-14). Later, the astonishing resurrection of Jesus made salvation possible not only for Jews, but for all people in Christ (Gal 3:26-29). Ultimately, the New Testament makes it clear that God’s renewal will encompass the entire Creation:

For God was pleased to have all his fullness dwell in him, and through him to reconcile to himself all things, whether things on earth or things in heaven, by making peace through his blood, shed on the cross. (Colossians 1:19-20)

With all wisdom and understanding, he made known to us the mystery of his will according to his good pleasure, which he purposed in Christ, to be put into effect when the times reach their fulfillment—to bring unity to all things in heaven and on earth under Christ. (Ephesians 1:8-10)

Christians are accustomed to thinking of the death of Christ in regard to humans, but our culture rarely acknowledges God plan for the redemption of His entire creation. This is partly attributable to the fact that discussions of creation and origins are often separated from the topic of salvation.¹⁷ In doing so we tend to marginalize Jesus as we argue about Genesis. Rather than fall into this trap, if we view nature through a theology of the cross, we will see Christ as both the alpha and the omega point in discussions of life’s history and life’s future. With this perspective, we are more apt to sense our solidarity with the rest of creation as we wait in eager anticipation of a glorious future:

The creation waits in eager expectation for the children of God to be revealed. For the creation was subjected to frustration, not by its own choice, but by the will of the one who subjected it, in hope that the creation itself will be liberated from its bondage to decay and brought into the freedom and glory of the children of God. (Romans 8:19-21)

Conclusion

As part of the Church’s conversation about the problem of natural evil, this essay is meant to be a brief introduction to a “theology of the cross”. One can explore this concept in greater detail in Murphy’s book The Cosmos in the Light of the Cross. While there is a lot more to be said, let me conclude with the following observation: though evolution may not be compatible with some interpretations of Christianity, evolutionary creation is certainly compatible with the crucified Christ and the theology of the cross. In the person of Jesus, God suffers with the world and ultimately redeems it. As George Murphy puts in, “The world's pains are God's stigmata.”¹⁸

Explore this Topic Further

• Miller, Keith. “And God saw that it was good”: Death and Pain in the Created Order. BioLogos series
• Murphy, George L. The Cosmos in the Light of the Cross. Harrisburg, PA: Trinity Press, 2003.
• Murphy, George L. “Cross, Evolution, and Theodicy: Telling It Like It Is”. In The Evolution of Evil. Edited by G. Bennett, M.J. Hewlett, T. Peters, and R.J. Russell. Göttingen: Vandenhoeck & Ruprecht, 2008.
• Southgate, Christopher. The Groaning of Creation: God, Evolution, and the Problem of Evil. Louisville, KY: Westminister John Knox Press, 2008.

Notes

1. Murphy, George L. The Cosmos in the Light of the Cross. Harrisburg, PA: Trinity Press, 2003.
2. Murphy, p34
3. “CRUX Sola Est Nostra Theologia,” in D. Martin Luthers Werke, Kritische Gesammtausgabe (Weimar: Hermann Boehlau, 1892), 5:172. The captitalization is in the original. Cited in Murphy, p26.
4. Murphy, p108
5. Murphy, p4
6. Murphy, p43
7. Murphy, p3
8. Some Christians ascribe animal death to some combination of Adam’s fall and Noah’s flood, but this does not resolve the problem that the animals are still suffering and dying through no fault of their own. See Keith Miller’s BioLogos series Death and Pain in the Created Order for the limitations inherent in a fall-based theodicy.
9. Rudwick, Martin. The meaning of fossils: Episodes in the history of paleontology. Chicago, University of Chicago Press, 1985.
10. See Jeff Schloss’ BioLogos essay Evolution, Creation, and the Sting of Death
11. Murphy, p63
12. Murphy, p122
13. Murphy, p27
14. This example is drawn from an evangelical outreach event held by a Christian student group in Innsbruck, Austria. On campus one day, they started conversations with their classmates by asking the question, “Would you wear an electric chair on your neck?”
15. Murphy, George L. The Trademark of God: A Christian Course in Creation, Evolution, and Salvation. Wilton, Conn.: Morehouse-Barlow, 1986.
16. Murphy, Cosmos in the Light of the Cross, p 31-32.
17. Murphy, p35
18. Murphy, p87

In Part 2 of this series, we concluded by noting that, as Christian geologists willing to consider the possibility, we find no compelling evidence that the earth’s geological features can be explained by a global Flood. Here we consider three lines of evidence: global salt deposits, the order of deposition of sediment layers in the Grand Canyon, and the sequence of fossils in geological strata.

Salt Deposits

There are many places around the earth with layers of salt, some thousands of feet in thickness. Just off the southern coast of the United States in the Gulf of Mexico, thick salt deposits sit beneath thousands of feet of sediment (Fig. 1). These deposits lie within the layers that are said to have been deposited by the Flood.

We understand how salt beds form. At locations such as the Bonneville Salt Flats of Utah, or at the Dead Sea at the border of Israel and Jordan, salt is actively forming. Salt beds form when water is evaporated. During evaporation, the concentration of dissolved ions increases until the water cannot hold the salt in solution anymore and mineral salt begins to form. If a presently unknown or poorly understood process could produce salt without evaporation, as argued by young-earth advocates¹, it would quickly dissolve as soon as it came into contact with flood water, just as the salt from your saltshaker rapidly dissolves when added to water or moist food.

One might argue that the waters from the Flood could have evaporated to leave behind the salt deposits we see today, but there is a serious problem. The thousands of feet of sediment on top of the salt is also said to be from the Flood, meaning the flood waters cannot have evaporated to produce the salt and still be present and violent enough to transport thousands of feet of sediment to the same location. In other words, a single flood cannot be called upon to explain both the salt and the overlying sediment. For those who wish to argue that natural processes could have been vastly different during the Flood, there are at least two replies. First, under such a scenario, there is no point in Flood Geology studies any more than in normal studies, for nothing could be gained by the study of unknowable processes. A more important question, however, would be to ask why God would alter natural processes just to make Flood sediments look like they are not flood sediments. What would the purpose be? (We will revisit this thought later.)

Grand Canyon: Order of Deposition

The Grand Canyon is made up of a sequence of layers that defies any reasonable attempt to explain by a single flood. The alternating layers of limestone, sandstone and shale each form in unique environments. If these deposits were formed at different times under various sea-level stages, it is quite simple to explain the different grain sizes and rock types as a function of depth and distance from the shore line. If explained with a single catastrophic flood that abided by God’s natural laws of physics and chemistry, logic must be stretched beyond the breaking point.

As a very simple observation, consider instructions given in virtually every gardening book. A good soil will have a mix of sand, silt and clay. To determine the quality of your soil, you take a handful or two, put it in a clear container, add water and shake it up. When you stop shaking, the coarse grained material will settle out first resulting in a sequence of layers: sand on the bottom, then silt, then clay. You can readily see how much of each you have by the thickness of each layer.

This is informative of what we see in flood deposits. As moving flood waters slow down, finer and finer grained sediment settles out resulting in a “fining upward” sequence. If most of the Grand Canyon layers were laid down by the Flood, then we should see the same thing – a “fining upward” sequence. Instead, we see a series of alternating layers of fine and coarse grained material, with smaller-scale alternating layers within the larger ones (Fig. 2). Increasing the violence of a flood does nothing to negate the standard order of deposition. Repeated surging of flood waters across the surface likewise offers little explanatory power; in this case we might expect successive layers, each with their own “fining upward” sequence, but such is not what is observed. Further, the Grand Canyon includes multiple layers of limestone, which are never found in flood deposits of any magnitude. Even in floods as massive as one thought to have catastrophically deluged the once dry Mediterranean Sea basin with thousands of feet of water – limestone beds are conspicuously absent.

Fossil Sequence

If a massive flood were responsible for the fossil record, what would we expect to see? If the Flood was violent enough to rip chunks of rock up from the earth and move entire continents (standard Young Earth claims)², then it should be obvious that life forms from every imaginable niche would be tumbled and mixed together (Fig. 3a). We should find numerous examples of mammoths mixed with triceratops, and pterodactyls mixed with sparrows. Ferns and meadow flowers should be found in the same deposits, along with trilobites and whales. Further, we should find all major life forms still living today, for Genesis 7:8-9 is clear in stating that all terrestrial animals were preserved on the ark.

What we actually observe is far different (Fig. 3b). There is an orderly sequence where trilobites only occur in very old rocks, dinosaurs in later beds, and mammoths in still later layers. Organisms like flowers and ferns are present together in more recent deposits, but only ferns with no flowers are found in older deposits. Some readers will recognize this as an example from the “geologic column” and be tempted to discount it as a fabrication. For those thinking this way, consider what Henry Morris had to say in both editions of Scientific Creationism:

“Creationists do not question the general validity of the geologic column, however, at least as an indicator of the usual order of deposition of the fossils…”⁵

If we revisit the Grand Canyon for a moment, is it not striking that there is not a single dinosaur, mammoth or bird in the entire exposed sequence? Not one. To find these, you have to go to younger sediments found in deposits outside the canyon that have not been fully eroded away yet. How could such a lack of mixing be possible if the Flood was violent enough to move continents?

For more, be sure to read our FAQs How are the ages of the Earth and universe calculated? and What scientific evidence do we have about the first humans?, as well as our recent infographic How Do We Know the Earth is Old?.

Author's Note from Joy Walters

As I mentioned in my first post, I grew up skeptical of the whole idea of evolution. One contributor to my disbelief was the lengthy timescale for the “tree of life” that was presented with the theory. I would hear, for example, that dinosaurs lived hundreds of millions of years ago, but there was no explanation of why this was true; it was just given as a fact. No one explained the methods of dating, and so I thought biologists simply estimated the ages of species to fit their preconceived notions of how long it would take for one species to emerge from another. It also seemed like the ages were periodically revised and extended farther back in time, and I figured scientists needed to manipulate numbers to make evolution plausible. This, in my mind, made the theory both unbelievable and dismissible.

Once I learned about the techniques used to date fossils, I realized that my first impressions were wrong; the ancient ages of species are scientific determinations rather than scholarly conjectures. However, I have found in recent conversations that Christians remain skeptical of old ages and the evolutionary time scale. For this reason, I wanted the videocast to address the process of fossil dating (what the methods are and why they are accurate) while focusing on cases where hominid fossils were discovered and dated using these very methods. My hope is that Believers would be informed about the evidence for human evolution and its scientific grounding.

There are two opposite errors which need to be countered about the fossil record: 1) that it is so incomplete as to be of no value in interpreting patterns and trends in the history of life, and 2) that it is so good that we should expect a relatively complete record of the details of evolutionary transitions within all or most lineages.

What then is the quality of the fossil record? It can be confidently stated that only a very small fraction of the species that once lived on Earth have been preserved in the rock record and subsequently discovered and described by science.

There is an entire field of scientific research referred to as "taphonomy" -- literally, "the study of death." Taphonomic research includes investigating those processes active from the time of death of an organism until its final burial by sediment. These processes include decomposition, scavenging, mechanical destruction, transportation, and chemical dissolution and alteration. The ways in which the remains of organisms are subsequently mechanically and chemically altered after burial are also examined -- including the various processes of fossilization. Burial and "fossilization" of an organism's remains in no way guarantees its ultimate preservation as a fossil. Processes such as dissolution and recrystallization can remove all record of fossils from the rock. What we collect as fossils are thus the "lucky" organisms that have avoided the wide spectrum of destructive pre- and post-depositional processes arrayed against them.

Soft-bodied organisms, and organisms with non-mineralized skeletons have very little chance of preservation under most environmental conditions. Until the Cambrian nearly all organisms were soft-bodied, and even today the majority of species in marine communities are soft-bodied. The discovery of new soft-bodied fossil localities is always met with great enthusiasm. These localities typically turn up new species with unusual morphologies, and new higher taxa can be erected on the basis of a few specimens! Such localities are also erratically and widely spaced geographically and in geologic time.

Even those organisms with preservable hard parts are unlikely to be preserved under "normal" conditions. Studies of the fate of clam shells in shallow coastal waters reveal that shells are rapidly destroyed by scavenging, boring, chemical dissolution and breakage. Occasional burial during major storm events is one process that favors the incorporation of shells into the sedimentary record, and their ultimate preservation as fossils. Getting terrestrial vertebrate material into the fossil record is even more difficult. The terrestrial environment is a very destructive one: with decomposition and scavenging together with physical and chemical destruction by weathering.

The potential for fossil preservation varies dramatically from environment to environment. Preservation is enhanced under conditions that limit destructive physical and biological processes. Thus marine and fresh water environments with low oxygen levels, high salinities, or relatively high rates of sediment deposition favor preservation. Similarly, in some environments biochemical conditions can favor the early mineralization of skeletons and even soft tissues by a variety of compounds (eg. carbonate, silica, pyrite, and phosphate). The likelihood of preservation is thus highly variable. As a result, the fossil record is biased toward sampling the biota of certain types of environments, and against sampling the biota of others.

In addition to these preservational biases, the erosion, deformation and metamorphism of originally fossiliferous sedimentary rock have eliminated significant portions of the fossil record over geologic time. Furthermore, much of the fossil-bearing sedimentary record is hidden in the subsurface, or located in poorly accessible or little studied geographic areas. For these reasons, of those once-living species actually preserved in the fossil record, only a small portion have been discovered and described by science. However, there is also the promise of continued new and important discovery.

The forces arrayed against fossil preservation also guarantee that the earliest fossils known for a given animal group will always date to some time after that group first evolved. The fossil record always provides only minimum ages for the first appearance of organisms.

Because of the biases of the fossil record, the most abundant and geographically widespread species of hardpart-bearing organisms would tend to be best represented. Also, short-lived species that belonged to rapidly evolving lines of descent are less likely to be preserved than long-lived stable species. Because evolutionary change is probably most rapid within small isolated populations, a detailed species-by-species record of such evolutionary transitions is unlikely to be preserved. Furthermore, capturing such evolutionary events in the fossil record requires the fortuitous sampling of the particular geographic locality where the changes occurred.

Using the model of a branching tree of life, the expectation is for the preservation of isolated branches on an originally very bushy evolutionary tree. A few of these branches (lines of descent) would be fairly complete, while most are reconstructed with only very fragmentary evidence. As a result, the large-scale patterns of evolutionary history can generally be better discerned than the population-by-population or species-by-species transitions. Evolutionary trends over longer periods of time and across greater anatomical transitions can be followed by reconstructing the sequences in which anatomical features were acquired within an evolving branch of the tree of life.

Today's video is courtesy of filmmaker Ryan Pettey, director/editor of Satellite Pictures and features physicist Ard Louis.

In today's video, Oxford physicist Ard Louis discusses the famous debate between renowned evolutionary biologists Stephen Jay Gould and Simon Conway Morris. Gould believed (and wrote in his book Wonderful Life) that if the "tape" of evolution were rerun, the chance that anything like human intelligence would emerge is essentially zero. In other words, humanity is here through random accident. Gould pointed to the work of Morris and fellow scientists in their research of the Burgess Shale as evidence for this view.

However, Morris himself disagrees, pointing to what is called evolutionary convergence. As Morris notes, there are numerous examples of identical features evolving multiple times throughout the history of life independently. Morris believes that if the tape of life were replayed, we would see something like humans emerge. A Christian might say, it looks like we were planned.

Some Christians might find Simon Conway Morris' viewpoint, with its implicit teleology, more attractive. Others, perhaps motivated by a high view of providence, may find Gould's emphasis on contingency equally congenial to their faith. What do you think?

In our previous essay, we learned that a tree summarizing species relationships can be built using DNA information, and how we can use DNA as a “molecular clock” to date ancient events. Both of these methods have made specific predictions about the origin of whales. If evolution is true: whales are related to the even-toed hoofed mammals and should share common ancestors with them; transitional fossil forms dating from about 45 to 50 million years ago should be found which can be shown to be related to both the even-toed hoofed mammals and modern whales; whales are most closely related to modern hippos, and should share a common ancestor with them.

What other types of information might we be able to use to construct a phylogenetic tree (i.e. a family tree) of species relationships? It turns out that characteristics of body structure also can be used – for example, the presence or absence of certain bones, or the specific shapes of those bones. An advantage of using bony features is that they can be recovered from fossils, whereas DNA (with only certain limited exceptions) must come from living organisms.

We can also derive functional information from an examination of bony features. The various protrusions, bumps and knobs found on bones usually have important implications. For example, smooth rounded areas at the ends of bones allow them to fit together and move easily. The shapes of such surfaces determine which bone motions are “allowed” or “disallowed”. Consider, for example, the motion of the forearm against the upper arm at the elbow. This is a “hinge” joint, whose normal motion is defined by the shapes of the upper arm bone and one of the forearm bones, where they meet each other. You might normally exercise the action of this hinge joint when you pick up a cup of coffee, bring it to your mouth, then set it back down again. Let’s try to imagine another motion. For this exercise we first need to get our arm into the proper starting position. Place your arm at your side, bent at the elbow at a ninety degree angle, with your palm up. Now, while keeping your palm up, let’s attempt to move your arm only at the elbow (no shoulder motion – that’s cheating!). Now swing your forearm out to the side and attempt to end up with your fingers pointed directly away from your side. Most of you will not be able to do this. If you can, it’s because your shoulder is rotating in spite of yourself. This motion at the elbow is normally not allowed. Hence a careful analysis of bone shapes can allow us to infer how the bones were used. This in turn can assist us in the task of phylogenetic (evolutionary) classification of organisms. That is, we will have more confidence in the grouping together of animals in our tree diagram if corresponding bones are used functionally in the same way.

Therefore we would expect that we could use various bony features to help us examine the predictions generated by our previous look at different types of DNA data. Are there any bony features that are particularly relevant to the even-toed hoofed mammals? Well, it turns out that there are. These are mainly running animals, and there are several features of their ankle bones, which taken together define the “allowed” motions which make them efficient runners. If one takes the various ankle bones of a large group of mammals, examines them carefully to note their shapes, scores that information into a table, then uses a computer program to build a phylogenetic tree, it turns out that all the even-toed hoofed mammals are placed together. So far, so good. But what about whales? Well, now we have an obvious problem. Modern whales are very specialized, - they have flippers which correspond to the forelimbs, and they have almost no hind limbs! I say “almost” because they do have small pelvic bones, which are not attached to the rest of their skeletons. But they certainly have no ankles. This is where the fossils should come in – if evolution is true, we should expect to be able to identify transitional fossils which are ancestral to whales which contain the characteristic ankle bony features of the even-toed hoofed mammals.

Now let’s look at bony features from the whale perspective. We have already mentioned the almost complete loss of hind limbs, and the presence of forelimbs modified into flippers. In addition, as air breathers, whales have a blowhole at the top of their skull. And as powerful swimmers, which use a large tail fluke in vertical motions, whales have enormous sets of muscles which attach to enlarged projections from their vertebral column. So if evolution is true, we should begin to see fossil forms which manifest changes in bony features which correspond to the gradual accumulation of these whale-like characteristics. However, we still need more, because these various bony features all would be expected to occur in largely or exclusively aquatic forms. We might expect this to correspond to the later stages of a transition from terrestrial even-toed hoofed mammals. But what about the earlier stages? It would be very helpful if we had some “defining” characteristic of whales, similar to the ankle structure of even-toed hoofed mammals.

It turns out that the structure of the bones of the skull and ear apparatus of whales are highly modified to allow efficient hearing underwater. The mechanical aspects of efficiently receiving sound through water are somewhat different than receiving sound travelling through air. If evolution is true, we should expect to be able to find key transitional fossil forms with a progressive series of modifications of the skull and ear bones, features which would not be found in any other mammals.

Now that we know what we should expect to see, if evolution is true, let’s look at what has actually been found in the fossil record. Over the last fifteen years or so, a series of fossils, many of them discovered in the Indian subcontinent, have fulfilled nearly all of our predictions.^1,2 Let’s look at the figure below (Figure 1), reproduced from a recent popular book on evolution.³ This shows a series of fossils, arranged in approximate chronological order, with a modern whale at the top. How old are these fossil forms? The entire fossil progression illustrated occurs from a little over 50 million years ago to about 40 million years ago. So a remarkable alteration in general body form occurred in a little over 10 million years. This time frame agrees well with the previous prediction from the DNA “clock” that we discussed in our previous essay. Second, the general change in body shape corresponds to what we predicted in our discussion of whale bony features above. That is, there is a gradual elongation and streamlining, there is a modification of the forelimb into flippers and progressive reduction of the hindlimb, the nostrils for breathing move toward the top of the skull to form a blowhole (not obvious from the diagram), and the vertebrae develop enlarged projections to support the attachment of swimming muscles.

Figure 1: Skeletons and Body Forms of Modern Whales and Fossil Ancestors
The reconstructed skeletons (black) from modern whales (top) and various ancestral skeletal forms (series below) are in chronological order (from Pakicetus up). Indohyus is an extinct whale “cousin”. Relative body size, to scale, is indicated by the gray shapes at the right of each animal.

There is probably little question that the last fossil species in the figure (Durodon) is well on the way to becoming a modern whale. However, it might be argued by a skeptic that the earlier species (like Rhodocetus, Ambulocetus, or Pakicetus), despite the “cetus” (whale) part of their names, are not so obviously “whale-like” that they deserve to be considered as fossil whale ancestors. However, remember the characteristic whale skull modifications for hearing? It has been shown very clearly that throughout this series of fossil species, the various bony changes necessary to support efficient hearing in water were being acquired in a stepwise fashion. Organisms earlier in the sequence had skeletal characteristics consistent with them being able to hear well in both air (using the “classic” mammalian hearing apparatus), and newly acquired changes to also allow better hearing in water. Later organisms in the sequence become increasingly specialized for hearing in water only.⁴

What of the earliest fossil shown in this diagram –Pakicetus? Careful examination shows that it has the features we would predict for an early whale ancestor. It has the ankle bone characteristics of the even-toed hoofed mammals (in fact these features are also found in several of the later fossil forms as well, ensuring their continuity). This confirms one of the predictions made by the DNA evidence we discussed earlier. Furthermore, it has some of the modifications of the skull bones necessary for more efficient underwater hearing, which were previously documented only for modern whales and their later (more obvious) ancestors.⁴ These features are also shared with the “whale cousin” Indohyus.⁵ Preservation of more of the skeleton of this latter species has allowed detailed analysis indicating characteristics likely shared with whale ancestors. Indohyus was probably a wading animal, which spent much of its time in the water. It appears to have fed mostly on land, so it is suggested that resort to the water was made to escape predators.⁵

Finally, we need to look back at the last prediction from our previous DNA evidence, namely that modern whales are most closely related to hippos. If evolution is true, we should expect to find fossil forms linking these two modern groups. This has proven to be a tougher nut to crack, mainly because the ancestral whales first appear about 50 million years ago in what is now south Asia, and the hippo family first appears about 15 million years ago, in Africa. The most recent tree diagram, produced by using a combination of skeletal features and DNA data, still supports this family connection, as shown by the following figure (Figure 2).⁶

Figure 2: Phylogenetic Tree Showing the Relationship of Modern Whales to Living and Extinct Even-Toed Hoofed Mammals
This tree is based on both bony features and DNA data. The organisms presented in blue are semi-aquatic or aquatic forms. Organisms shown in green are terrestrial even-toed hoofed mammals (Artiodactyls). In black is shown a member of the odd-toed hoofed mammals. In red is an extinct fossil ancestor group. (This figure is adapted from Fig 1a in Reference 6).

The blue lines in the diagram show species in which the skeleton is specially thickened, and the bone structure more dense. This is an adaptation which allows wading animals (like modern hippos and the fossil Indohyus) to be good “bottom-walkers” (it prevents them from floating due to lighter body tissues), and allows fully marine organisms (like modern whales) to have “neutral buoyancy” (so they don’t always tend to pop up to the water surface, like a cork). There has also been progress in clarifying the relationships between fossil ancestors of hippos and those of modern whales. A recent study of hippo evolution, based only on skeletal characteristics, has conclusively shown that the hippo family are descended from an extinct group of fossil Artiodactyls, known to go back more than 40 million years, and whose fossils are from southern Asia. Furthermore, this study produced a phylogenetic tree predicting that this extinct hippo ancestor group also shared a common ancestor with the fossil whales.⁷ Thus the investigation of hippo origins is independently leading us back toward the origin of whales. However, in this study the statistical support for predicted common ancestor of the ancient hippo group and the ancient whale group is not as strong as scientists would like to consider this “case closed”. What is necessary is more fossils, of the appropriate age in order to complete the story of hippo evolution. We still need that to fill in the details of the predicted relationship of hippos to modern whales.

Thus the “Whales’ Tale” is not yet complete. It is a story of scientific discovery in progress. As we finish, let’s briefly summarize what we have found out. Different types of DNA evidence agree that modern whales are most closely related to the even-toed hoofed mammals, despite the obvious great changes in limb anatomy of the modern whales. This prediction has been amply confirmed by the fossil record. The DNA sequence evidence predicted a time frame during which critical early events in evolution of whale ancestors should occur. This prediction has also been amply confirmed. Finally, DNA evidence predicts that modern whales are most closely related to hippos. There is some fossil evidence supporting a predicted common ancestor, but more data is needed. A final caution to possible sceptics – this state of “unfinished business” is precisely how the scientific process works. There is no “crisis”. There is no indication that evolution is not true. There is simply the ongoing work of mapping out of various lines of evidence. A scientific conclusion is considered well supported if “all roads lead to Rome”. In the case of whale evolution it might be prudent to say that the evidence has not quite converged in Rome yet, but that we are now in the suburbs. That is precisely what makes science interesting and fun. Stay tuned!

The next blog in this series can be found here.

References:

1: Thewissen J.G.M., Williams E.M., Roe L.J. and Hussain S.T. 2001. Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls. Nature. 413: 277-281.

2: Gingerich P.D., ul Haq M., Zalmout I.S., Khan I.H., Malkani M.S. 2001. Origin of Whales from Early Artiodactyls: Hands and Feet of Eocene Protocetidae from Pakistan. Science. 293:2239-2242.

3: Coyne, J.A. 2009. Why Evolution is True. Viking Penguin, New York. Pg 50.

4: Numella S., Thewissen J.G.M., Bajpai S.,Hussain T., Kumar K. 2007. Sound Transmission in Archaic and Modern Whales: Anatomical Adaptations for Underwater Hearing. The Anatomical Record. 290:716-733.

5: Thewissen J.G.M., Cooper L.N., Clementz M.T., Bajpai S., Tiwari B.N. 2007. Whales originated from aquatic artiodactyls in the Eocene epoch of India. Nature. 450:1190-1194.

6: Geisler J.H. and Theodor J.M. 2009. Hippopotamus and whale phylogeny. Nature. 458:E1-E4.

7: Boisserie J.-R., Lihoreau F., Brunet M. 2005. The position of Hippopotamidae within Cetartiodactyla. Proceedings of the National Academy of Sciences U.S.A. 102(5):1537-1541.

In the rest of the series, we will look closely at the evolutionary explanation for the blueprint. That explanation postulates that tetrapod limbs arose during a particular era in life's history, and that they arose as modifications of the fins of fish. And the evolutionary explanation isn't just an interesting idea. It's a comprehensive explanation – it helps us to understand bones, fossils, genes, chemical signaling systems. It provides a coherent framework for understanding why limbs are the way they are, and how they got that way.

Comparing the anatomy of tetrapod vertebrates and fish

The idea that tetrapods arose from fish is not new; E.D. Cope proposed in 1892 that tetrapods descended from lobe-finned fish. (Modern lobe-finned fishes include coelacanths and lungfish, and comprise one of two divisions of the bony fishes. The other division, the ray-finned fishes, includes most familiar kinds of fish.) In the early days, biologists inferred ancestral relationships between species largely through comparative anatomy and embryology: they would carefully classify organisms according to their structure (including their structure during development) and look for relationships that generated nested hierarchies. A simple nested hierarchy goes something like this: 1) animals with backbones; 2) animals with backbones and limbs; and 3) animals with backbones, limbs, and hair. Animals in group 3 also belong to groups 1 and 2, while animals in group 2 also belong to group 1, and so the groups together define a nested hierarchy. Such studies alone could have led some scientists to infer an ancestral relationship between fish and tetrapods, and perhaps those studies did convince them. But then there were fossils of various types of fish and other vertebrates, many long extinct. That fossil record was relatively sketchy in 1892, but it nevertheless led Cope and others to conclude that certain fish had given rise to tetrapods at a particular time in natural history.

The fossil record shows a fish-to-tetrapod transition

Here are some basic findings from the fossil record that suggest a fish-to-tetrapod transition and that have been known for decades:

• Fish, including fish with bones, lived on the earth before tetrapods appeared. Specifically, fossils of bony fish first appear in rocks from about 420 million years ago.

• Tetrapods appear in the fossil record at a particular point in history and then persist and diversify in subsequent eons. Their arrival was long thought to have occurred about 365 million years ago, although some recent findings have challenged that hypothesis.

• Tetrapods that still had some fishy features were prowling the planet 365 million years ago.

• Lobe-finned fish that were starting to look more like tetrapods were eating other fish about 385 million years ago.

Even many decades ago, there were hints that something interesting happened between 400 million years ago and 365 million years ago. Let's take a close look at the ancient animals that suggest the fish-to-tetrapod transition.

Ancient animals

The most primitive known tetrapods for which we have skeletal remains lived 365 million years ago. They were undeniably tetrapods, but there was definitely something fishy about them. (Heh heh.) One of the most famous of these creatures is Acanthostega, discovered in 1987 by British paleontologist Jennifer Clack and pictured below. Acanthostega is a card-carrying tetrapod, with fingers and toes. But it has a fish tail, with fin rays. Another well-known primitive tetrapod is Ichthyostega, which lived around the same time as Acanthostega. Like Acanthostega, it is a true tetrapod, but has several odd fish-like structural features. For example, its skull is more fish-like than that of Acanthostega. In summary, both Acanthostega and Ichthyostega already used the limb blueprint, even though both also had some fish-like anatomical characteristics. Their presence 365 million years ago shows that tetrapods must be at least that old, and their mixture of anatomical features suggests that the transition happened not long before that.

And what about the lobe-finned fish that looked a bit tetrapod-ish? That animal is Panderichthys, described as “vaguely crocodile-shaped” with skeletal features that were tetrapod-like. Specifically, this ancient fish had tetrapod-like “shoulders,” and recent analysis found some finger-like bones at the ends of the fins. The creature also had a breathing hole on the top of its head. These fish lived around 385 million years ago.

The hunt for the earliest tetrapods

Taken together, these observations suggested that the fish-to-tetrapod transition occurred between 385 and 365 million years ago. Eager to see what that transition looked like, scientists began to look for 375 million-year-old rocks in which they might find animals at the beginning of tetrapod-hood. They wanted to catch evolution in the act.

Let's stop and think about this, because it's cool and because it's important to note the extent to which evolutionary biology is hypothesis-driven. Critics of evolution sometimes portray the theory as an untestable historical conjecture, depicting it as fundamentally different from experimental science in the lab. But the hunt for the earliest tetrapods was an effort to test a hypothesis that had generated a prediction. Based on the hypothesis that lobe-finned fish were ancestors of tetrapods, scientists predicted that intermediate animals, “fishapods,” would be found in the gap between Panderichthys and Acanthostega. To evaluate the prediction, all they needed to do was find some suitable 375 million-year-old rocks.

Neil Shubin describes that search in the first chapter of Your Inner Fish. He and his colleagues found suitable rocks in the islands of the Arctic: the right age, nicely exposed (by erosion), and representative of the kind of environment that their quarry would frequent – freshwater streams. They made their biggest discovery on their last trip (“a do-or-die situation”) in 2004. That discovery was Tiktaalik roseae, the “fishapod.”

The “fishapod”

Tiktaalik roseae is one of the most extraordinary fossil intermediates ever described, and its public debut in 2006 was front-page news. An artist's conception of the animal is pictured below.

There are several aspects of the anatomy of Tiktaalik that earn it the title “fishapod.” Like a good fish, it had scales and webbed fins. Like a tetrapod (more specifically, like a crocodile), it had a flat head, with eyes on the top of the head, and it had a neck. But what about those fins? Or are they limbs? Remarkably, the answer seems to be, “both.”

The fins of Tiktaalik were part fish fin, part tetrapod limb. On the outside, they looked like fins, with webbing. On the inside, though, these fins were clearly tetrapod-like. Amazingly, the fins of Tiktaalik were built using a primitive version of the limb blueprint: one bone, two bones, blobs, digits. As Shubin writes in Your Inner Fish, “We had a fish with a wrist.” (The Tiktaalik roseae web site at the University of Chicago is a great source for images and more information.)

Let's address three questions about Tiktaalik that might have occurred to you. First, why might animals like Tiktaalik have developed tetrapod-like fins? Shubin and his colleagues suggest that these limb-like fins may have been useful for doing “push-ups” in the shallow water. (Like Panderichthys, Tiktaalik had a breathing hole on top of its head and was clearly adapted for living and moving in shallow water.) Second, is Tiktaalik an ancestor of all tetrapods? No, not necessarily. What Tiktaalik shows us is that animals were developing tetrapod features in the context of fish bodies, and Tiktaalik shows us the context (shallow water) in which this likely occurred. But that doesn't mean that our lineage arose from Tiktaalik itself. Finally, is Tiktaalik now the oldest tetrapod? No, apparently not. For one thing, Tiktaalik is truly transitional, and probably therefore not worthy of full tetrapod membership. But more notably, data published in 2010 show that tetrapods are a lot older than was thought at the time of Tiktaalik's discovery. The new findings show footprints that are unmistakably those of a tetrapod, in rocks about 395 million years old. Surprisingly, then, tetrapods were already on their way long before Neil Shubin's specimen lived. Tiktaalik is truly a fish/tetrapod intermediate, which was living at the same time as animals that were fully tetrapods. A simple story of succession, in which intermediates disappear and are replaced by less intermediate types, seems to be an oversimplification.

In conclusion, the fossil record provides evidence that the fins of fish and the limbs of tetrapods are related by ancestry: limbs seem to be modified versions of fins. What other evidence supports this proposal? In the next post, we will turn to developmental biology, and explore the meaning of the term 'homology.'

Further reading:

Neil Shubin (2009) Your Inner Fish: A Journey Into the 3.5-Billion-Year History of the Human Body. New York: Vintage Books.

Carl Zimmer (2006) A Fin is a Limb is a Wing: How Evolution Fashioned its Masterworks. Online at NationalGeographic.com.

Tiktaalik roseae website at the University of Chicago.

Jennifer Clack's website at the University of Cambridge.

Images are from Wikipedia.

Introduction: What’s all the fuss?

The most fundamental claim of biological evolution is that all living organisms represent the outer tips of a diversifying, upward-branching tree of life (click image to enlarge). The “tree of life” is an extremely powerful metaphor that captures the essence of evolution. Like the branches of a tree, as we trace individual lines of descent (lineages) back into the past (down the tree) they converge with other lineages toward their common ancestors. Similarly, these ancient lineages themselves converge with others back in time. Thus, all organisms, both living and extinct, are ultimately connected by an unbroken chain of descent with modification to a common ancestral trunk among single-celled organisms in the distant past.

This tree metaphor applies as much to the emergence of the first representatives of the major groups of living invertebrates (such as annelids, snails, or arthropods) as it does to the first appearance and diversification of dinosaurs, birds, or mammals. This early diversification of invertebrates apparently occurred around the time of the Precambrian/Cambrian boundary over a time interval of a few tens of millions of years. This period of rapid evolutionary diversification has been called the “Cambrian Explosion.”

The Cambrian explosion has been the focus of extensive scientific study, discussion, and debate for decades, and is increasingly receiving attention in the popular media. It has also received considerable recent attention by evolution critics as posing challenges to evolution. These critics argue that the expected transitions between major invertebrate groups (phyla) are absent, and that the suddenness of their appearance in the fossil record demonstrates that evolutionary explanations are not viable.

What are some of the arguments of the evolution critics? John Morris of the ICR writes:

“If evolution is correct, the first life was quite simple, evolving more complexity over time. Yet the Cambrian Explosion of Life has revealed life's complexity from the start, giving evolution a black eye. The vast array of complex life that appears in the lowest (or oldest) stratigraphic layer of rock, with no apparent ancestors, goes hard against evolutionary dogma. Evolution's desperate attempt to fill this gap with more simple ancestral fossils has added more injury. .... Think of the magnitude of this problem from an evolutionary perspective. Many and varied forms of complex multi-celled life suddenly sprang into existence without any trace of less complex predecessors. There are numerous single-celled forms at lower stratigraphic levels, but these offer scant help in solving the mystery. Not one basic type or phyla of marine invertebrate is supported by an ancestral line between single-celled life and the participants in the Cambrian Explosion, nor are the basic phyla related to one another. How did evolution ever get started?”¹

Intelligent design advocate Stephen Meyer and others have written:

“To say that the fauna of the Cambrian period appeared in a geologically sudden manner also implies the absence of clear transitional intermediates connecting the complex Cambrian animals with those simpler living forms found in lower strata. Indeed, in almost all cases, the body plans and structures present in Cambrian period animals have no clear morphological antecedents in earlier strata.²

And:

“A third feature of the Cambrian explosion (as well as the subsequent fossil record) bears mentioning. The major body plans that arise in the Cambrian period exhibit considerable morphological isolation from one another (or “disparity”) and then subsequent “stasis.” Though all Cambrian and subsequent animals fall clearly within one of a limited number of basic body plans, each of these body plans exhibits clear morphological differences (and thus disparity) from the others. The animal body plans (as represented in the fossil record) do not grade imperceptibly one into another, either at a specific time in geological history or over the course of geological history. Instead, the body plans of the animals characterizing the separate phyla maintain their distinctive morphological and organizational features and thus their isolation from one another, over time.”³

Are these critiques warranted? To what extent is the Cambrian explosion really problematic for the evolutionary picture of an unbroken tree of life extending back to the earliest life on Earth?

Geologic Time Scales: How big was the bang?

The relative rapidity of the diversification of invertebrates during the Cambrian “explosion” is set against the backdrop of the Earth’s geologic and biologic history. Geologic time is unfamiliar to most people, and its shear vastness is difficult to grasp.

Two lines of evidence impact our understanding of the duration of the animal diversification that led to the appearance of the major groups of living invertebrates. The first is the dating of critical strata within the geological timeline such as the Precambrian-Cambrian boundary and various important fossil-bearing horizons. The second is the time of appearance of the first widely recognized fossil representatives of the major living groups (phyla) of invertebrate animals. The latter is in considerable flux as new fossil discoveries are made.

Originally, the base of the Cambrian had been set at the earliest appearance of organisms with mineralized skeletons - particularly trilobites. However, a diverse collection of tiny mineralized plates, tubes and scales was discovered to lie below the earliest trilobites.⁴ This interval of “small shelly fossils” was designated the Tommotian. Because of the presence of even earlier tiny mineralized tubes and simple burrows, there was no internationally accepted definition for the boundary until 1994. At that time, the base of the Cambrian was placed at the first appearance of a particular collection of small fossil burrows characterized by Treptichnus pedum.

Until the early 1990's the age of the Precambrian-Cambrian boundary was not tightly constrained, and was estimated to be about 575 million years ago. However, in 1993 new radiometric dates from close to the accepted Precambrian-Cambrian boundary revealed that it was significantly younger -- about 544 million years.⁵ A more precise date of 542 ± 0.3 million years has recently been formally accepted by the International Commission on Stratigraphy. The basis for this date was the discovery that a sharp worldwide fall (or negative spike) in the abundance of the isotope carbon-13 was coincident with the Cambrian boundary as previously defined. In Oman, this isotopic marker also coincides with a volcanic ash layer that yielded the 542 million year date using uranium/lead radiometric methods.⁶ This horizon also marks the last occurrence of several fossils characteristic of the underling late Precambrian Ediacaran Period.⁷ Such extinction events are commonly used to subdivide the geologic time scale.

The earliest diverse fossil invertebrate communities of the Cambrian are represented by the Chengjiang, in China. These deposits are dated at 525-520 million years. The famous Burgess Shale is considerably younger, dating at about 505 million years, and the end of the Cambrian Period is set at 490 million years. The Cambrian Period thus lasted for 52 million years. To put this in perspective, the time elapsed since the extinction of the dinosaurs at the end of the Cretaceous has been 65 million years. The Cambrian was a very long period of time.

If the Cambrian explosion is understood to comprise the time from the base of the Cambrian to the Chengjiang fossil beds, then this period of diversification in animal body plans appears to have lasted about 20 million years. However, not all living animal phyla with a fossil record first appear within this time window. The colonial skeleton-bearing bryozoans, (click for image) for example, are not known from the fossil record until the end of the Cambrian around 491 million years ago.⁸ More significantly, several living invertebrate phyla have a fossil record that extends into the late Neoproterozoic before the Cambrian. Sponges (click for image) have been recognized as early as 580 million years, cnidarians (click for image--the group includes jellyfish and anemones) are present among the Ediacaran animals at around 555 million years, and the stem groups (see discussion below) for some other phyla were also likely part of the Ediacaran communities.

Defining the Cambrian “explosion” is not as straightforward as it might seem. Although there was clearly a major burst of evolutionary innovation and diversification in the first 20 million years or so of the Cambrian, this was preceded by an extended period of about 40 million years during which metazoans (multicellular animals) arose and attained critical levels of anatomical complexity. The Ediacaran saw the appearance of organisms with the fundamental features that would characterize the later Cambrian organisms (such as three tissue layers, and bilaterally symmetric bodies with a mouth and anus), as well as the first representatives of modern phyla. The base of the Cambrian is not marked by a sharp dramatic appearance of living phyla without Precambrian roots. It is a subjectively defined point in a continuum. The Cambrian “explosion” appears to have had a “long fuse.”

Notes

1. Morris, J.D., 2008, The Burgess shale and complex life, Acts & Facts 37 (10): 13.
2. Meyer, S.C., M. Ross, P. Nelson, & P. Chien. 2003. The Cambrian explosion: biology's big bang. Pp. 323-402 in J. A. Campbell & S. C. Meyer, eds., Darwinism, Design and Public Education: Michigan State University Press, Lansing, p. 326.
3. Meyer, S.C., M. Ross, P. Nelson, & P. Chien. 2003. The Cambrian explosion: biology's big bang. Pp. 323-402 in J. A. Campbell & S. C. Meyer, eds., Darwinism, Design and Public Education: Michigan State University Press, Lansing, p. 333.
4. Rozanov, A.Y., 1984, “The Precambrian-Cambrian boundary in Siberia,” Episodes 7: 20-24. Rozanov, A.Y., and A.Y. Zhuravlev, 1992, “The Lower Cambrian fossil record of the Soviet Union,” IN J.H. Lipps and P.W. Signor (eds.), Origin and Early Evolution of the Metazoa: Plenum, New York, p.205-282.
5. Bowring, S.A,, J.P. Grotzinger, C.E. Isachsen, A.H. Knoll, S.M. Pelechaty, and P. Kolosov, 1993, “Calibrating rates of Early Cambrian evolution,” Science 261: 1293-1298.
6. Gradstein, F.M., J.G.Ogg, A.G. Smith, et. al., 2004. A Geologic Time Scale 2004. Cambridge University Press.
7. Amthor, J. E.; J.P. Grotzinger,; S. Schröder, S.A. Bowring, J. Ramezani, M.W. Martin, and A. Matter, 2003, "Extinction of Cloudina and Namacalathus at the Precambrian-Cambrian boundary in Oman". Geology 31: 431–434.
8. Landing, E., A. English,and J.D. Keppie, 2010, “Cambrian origin of all skeletonized metazoan phyla - Discovery of Earth’s oldest bryozoans (Upper Cambrian, southern Mexico),” Geology 38: 547-550.

The bones in the limbs of all land vertebrates are remarkably similar. Feel around your upper arm, you'll only find one bone in there -- the humerus. Now move to your lower arm. If you're thorough, you will find two bones -- the radius and the ulna. As you continue down to the wrist, there is a whole set of little bones that attach to the metacarpals of the hand and the phalanges of the fingers and thumb. If your dog is nearby, see if he will let you continue your experiment with him. His bones are arranged a little differently, but the setup is the same. This pattern is remarkably consistent in land vertebrates. Even marine mammals like whales have a humerus, then a radius and an ulna followed by smaller bones. Darwin noted this consistency in his book On the Origin of Species. He writes, "What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include similar bones, in the same relative positions?"

It would be fun to watch Darwin read a couple of pages from the book Your Inner Fish by paleontologist Neil Shubin. Shubin studies fossils of vertebrates that lived at the time of transition from water to land. As he began his project, Shubin knew that amphibian fossils were abundant in 365 million year old rock formations, but totally absent globally in formations older than 385 million years. So he and his colleague, Ted Daeschler, decided that if they were to find the intermediates between fish and amphibians, they would need to look for fossils in exposed rocks that were about 375 million years old. A simple search through a geology textbook told them only one rock formation would meet their criteria perfectly. This formation was in northern Canada and was formed by sedimentation in a river delta.

After a six-year-long hunt from 1998-2004, Shubin and his team unearthed a "fish" very different than anyone had ever seen before. They had isolated a transitional organism with many characteristics of the land animals that became abundant 10 million years later, but it also had many characteristics of fish that lived in the period just before.

The book is a fun read, and is perfect for all who truly seek to understand God's method of creation. Here's why Darwin would have loved the story though: Shubin predicted the characteristics of a transitional species, the age of the rock formation, the type of environment in which it would be found (a river delta) and the geographic location (northern Canada) likely to have these fossils. He and his team then went out and searched for six painstaking years -- scanning the landscape inch by inch -- until they found it. It was a transitional organism with a variety of interesting features such as a neck, which was new to vertebrate life, and scales, which were an old feature. It had a rib cage with new features that came to characterize land animals, and it had interesting limbs described in meticulous detail. Just like you and your dog, the organism had the same key bones in the right place: the humerus, the radius, the ulna, some of the bones of the wrist and even primitive bones of the foot.

As beautiful as this story is, it does not end there. Unlike Darwin, we now know how the specific bones of the limb are built in an embryo, and we know why the pattern is so similar among the various land vertebrates. There is a set of genes that produces signals in a developing embryo. The hox D family of genes becomes active at specific locations in a developing limb, and bone is made.

What this means is that one group of cells in the expanding "limb bud" gets the signal, "make bone here." Further toward the tip of the limb bud, two blocks of cells receive the same signal, "make bone here." These two blocks of cells respond by making the radius and ulna. A little bit later and further toward the tip of the limb bud, a second wave of expression of the hox D signal is active. "Make bone in this region too," the signal says. The reason we all have the same pattern is that the hox D signaling pattern is very similar in the limb buds of the embryos for all land vertebrates. It can be tweaked to give variation in structure, but the basic pattern is virtually always present.

So what about the fins of fish? If fins gave rise to limbs in evolutionary history, one might expect they would have hox D gene expression which resembles that of limbs but would be different in a fairly significant way. The shark provides the best example; it has bones at the base of its fin. And, sure enough, in a shark embryo there is a wave of expression of the hox D genes in fin development just like in land vertebrates. Sharks don't have tiny bones that might correspond to our wrist, hand and fingers, so there are significant differences in hox D gene expression at the tip. However, the signaling pathway was already present in the fins of fish. The signal that says, "make bone here," was expressed in fish at the exact same location where limbs appear in land vertebrates. The transition had already begun as fish developed fins. At the most significant level, fins and limbs share important features.

I have been a biologist for a long time, and I hope I never stop getting shivers in my spine when I think about the beauty of how we come to know things in biology. Biologists make predictions, then they go out into the field or the lab to see if their predictions hold up. When hundreds of predictions of this sort are fulfilled, a theory reaches the point where it becomes certain, at least on a broad level. And that is where we are with evolution.

In this space, our purpose is not so much to try to persuade as it is to explain why we are so certain God created this way. If you wish to join us as we explore the ramifications, we would love to have you with us in the coming days. If you think that all of biology has it wrong, that is your prerogative. You're still welcome to read and think along with us. A great place to start is the book by Neil Shubin--it is very accessible to the general audience regardless of science background. Once you've read that, irrespective of what you personally believe, you'll understand why most Christian biologists view evolution as being God's way of creating life's diversity. It is my prayer that you may even see why the beauty in all of this draws us to our knees in worship.

The Cambrian Explosion is often posed as a challenge for evolution because the sudden burst of change in the fossil record appears to be inconsistent with the more typical gradual pace of evolutionary change. However, although different in certain ways, there are other times of very rapid evolutionary change recorded in the fossil record -- often following times of major extinction. The Cambrian Explosion does present a number of challenging and important questions because it represents the time during which the main branches of the animal tree of life became established. It does not create a challenge to the fundamental correctness of the central thesis of evolution, the descent of all living species from a common ancestor. This important period in the history of life extended over millions of years, plenty of time for the evolution of these new body plans (phyla) to occur. Furthermore, the fossil record provides numerous examples of organisms that appear transitional between living phyla and their common ancestors. The ongoing research about the Cambrian period is an exciting opportunity to advance our understanding of how evolutionary processes work, and the environmental factors shaping them.

The major animal body plans that appeared in the Cambrian Explosion did not include the appearance of modern animal groups such as: starfish, crabs, insects, fish, lizards, birds and mammals. These animal groups all appeared at various times much later in the fossil record.³ The forms that appeared in the Cambrian Explosion were more primitive than these later groups, and many of them were soft-bodied organisms. However, they did include the basic features that define the major branches of the tree of life to which later life forms belong. For example, vertebrates are part of the Chordata group. The chordates are characterized by a nerve cord, gill pouches and a support rod called the notochord. In the Cambrian fauna, we first see fossils of soft-bodied creatures with these characteristics. However, the living groups of vertebrates appeared much later. It is also important to realize that many of the Cambrian organisms, although likely near the base of major branches of the tree of life, did not possess all of the defining characteristics of modern animal body plans. These defining characteristics appeared progressively over a much longer period of time.⁴

Interpretations of the “Cambrian Explosion”

Not all scientists accept the idea that the Cambrian Explosion represents an unusually rapid evolutionary transition. The fossil record is notoriously incomplete, particularly for small and soft-bodied forms. Some researchers argue that the apparent rapid diversification of body plans is an artifact of an increase in the rate of fossilization, due in part to the evolution of skeletons, which fossilize more effectively.⁵ Many of the early Cambrian animals possessed some type of hard mineralized structures (spines, spicules, plates, etc.). In many cases these, often very tiny, mineralized structures are all that are found as fossils. There were major changes in marine environments and chemistry from the late Precambrian into the Cambrian, and these also may have impacted the rise of mineralized skeletons among previously soft-bodied organisms. ⁶

Most scientists are persuaded that something significant happened at the dawn of the Cambrian era and view the Cambrian Explosion as an area of exciting and productive research. For example, scientists are now gaining a better understanding of what existed before the Cambrian Explosion as a result of new fossil discoveries. Recent discoveries are filling in the fossil record for the Precambrian fauna with soft-bodied organisms like those in the Ediacaran Assemblages found around the world.⁷ Late Precambrian fossil discoveries also now include representatives of sponges, cnidarians (the group that includes modern jellyfish, corals and anemones), mollusks and various wormlike groups. Some of the new fossil discoveries, in fact, appear to be more primitive precursors of the later Cambrian body plans. The discovery of such precursors shows that the Cambrian organisms did not appear from thin air.⁸ Further discoveries will no doubt reveal more clearly the relationship of Precambrian organisms with the creatures found in the Burgess Shale and Chengjiang deposits.⁹

Genomic studies provide further insights into the origins of the Cambrian Explosion. Although the genetic divergence of organisms would have preceded the recognition of new body plans in the fossil record, accumulating genomic data is broadly consistent with the fossil record.¹⁰ Both point to the rise of the bilateria (bilaterally symmetric invertebrate animals) in the latest Precambrian Ediacaran, and their ecological explosion in diversity in the Cambrian.

Unanswered Questions

The sudden change of the Cambrian Era was, in relative terms, not too sudden for the process of evolution. The changes during the Cambrian Era did not occur over decades, centuries, or even thousands of years; they occurred over millions of years—plenty of time for evolutionary change. However, for millions of years beforehand, body plans of animals had remained relatively constant. Not until this time period did a significant change occur. The remaining questions are: What triggered the Cambrian Explosion? And why did so much change occur at this time? Several different theories address the origin of the Cambrian Explosion, proposing that dramatic environmental changes must have opened up new niches for natural selection to operate upon. These proposals include the runaway glaciation theory,¹¹ which proposes that glaciers briefly covered much of the earth, and the resultant loss of habitat created bottlenecks where evolution could act more rapidly. Another theory suggests that a change in atmospheric oxygen led to this sudden burst in evolutionary changes.¹² Yet another proposal is that major changes in the seafloor, from algae mat-covered surfaces in the late Precambrian to soft muddy bottoms later in the Cambrian, had dramatic evolutionary and ecological impacts.¹³

The Cambrian Era Fossils, Providing Answers

While the causes of the Cambrian Explosion remain a topic of open and exciting debate, the continued fossil discoveries from the Cambrian and Precambrian Eras are bringing more clarity to the evolutionary puzzle. These fossils provide valuable insight, particularly for envisioning the common ancestors of diverse groups. For instance, both vertebrates (fish) and echinoderms (sea urchins, starfish) are part of the group called deuterostomes. Without fossil evidence, it is hard to envision what a common ancestor would look like for these very different creatures. The Cambrian fossils are filling in the picture.¹⁴

Organisms have changed significantly over time. In rocks more than 1 billion years old, only fossils of single-celled organisms are found. Moving to rocks that are about 550 million years old, fossils of simple, multicellular animals can be found. At 500 million years ago, ancient fish without jawbones surface; and at 400 million years ago, fish with jaws are found. Gradually, new animals appear: amphibians at 350 million years ago, reptiles at 300 million years ago, mammals at 230 million years ago, and birds at 150 million years ago.¹ As the rocks become more and more recent, the fossils look increasingly like the animals we observe today.

The Transition to Land: Sea Creatures to Land Animals

Fossils of land animals, or tetrapods, first appear in rocks that are about 370 million years old. In older rocks, only sea creatures are found. But in 1998, scientists found a fossilized fin, 370 million years old, with eight digits similar to the five fingers humans have on their hands, as shown in Figure 1. However, the fin was undoubtedly that of a fish, which means this fossil is strong evidence of a transitional form.

Figure 1:An Illustration of the fossilized fin found in 1998. Its resemblance to a tetrapod is an indication of gradual evolutionary change from sea creatures to land animals. Source: Image is used by permission from Falk, Coming to Peace, 113.

One of the great success stories in the examination of the fossil record was the finding of a near-perfect fossilized transition between a vertebrate adapted for water and one adapted for land. Evolutionary biologist Neal Shubin set out to find a more complete transitional specimen than the 1998 fin. He determined the exact age of rock that he expected would yield a transitional land/water animal, and then he and his team spent four summers in the Arctic scouring rocks of that age to find one. The results (see Figure 2 below) were spectacular.²

From Reptiles to Mammals

Mammals first appeared in the fossil record about 230 million years ago, nearly 70 million years after reptiles first appeared. One group of reptiles, the cynodonts, first appeared about 260 million years ago and became increasingly mammal-like in more recent fossils—circa 245 million years ago. This change can be seen most clearly in the bone structure of the ear, as illustrated in Figure 3.

Figure 3: As shown in the image above, transitional fossils of cynodonts had two jaw hinges. These fossils date from a time when the dentary and squamosal bones were beginning to take over the role of jaw hinge (hinge #2). This allowed the articular and quadrate bones to evolve into the second and third bones of the mammalian ear, as shown on the right. Source: Image used by permission from Falk, Coming to Peace, 119. Originally from F. H. Pough, J. B. Heiser, and W. N. McFarland, Vertebrate Life, 4th ed. (Upper Saddle River, NJ: Prentice Hall, 1996), 607.

Scientists found a species of cynodonts, dating to just before the emergence of mammals, that had a double jaw hinge like that of a mammal. A pair of bones found in even earlier cynodont fossils seems to have transitioned slowly into the ear. No other fossils have been found that share a similar structure to the transitional cynodonts and date back before the time of mammals. Likewise, soon after mammals appeared, these cynodonts became extinct. This timing implies that the cynodont fossils record the transition from reptiles to mammals.³

Transitional Forms: Few and Far Between

Transitional forms occur just when one might expect to see a change from one body type to another. However, a common objection is that few transitional fossils have been discovered; thus many lineages cannot be traced smoothly.

There are several reason for these gaps in the fossil record. First, fossilization is a very rare event. Plus, transitional species tend to appear in small populations, where rapid changes in the environment can provide a stronger evolutionary drive. Finally, because fossilization itself is a rare event, smaller populations are sure to produce fewer fossils. The fact that transitional species have been found at all is remarkable, and it offers further support of gradual, evolutionary change.