The Cambrian “Explosion”, Transitional Forms, and the Tree of Life
This is part one in a series by Keith Miller. It is an updated and extension of Miller and Campbell's 2003 essay “The ‘Cambrian explosion’: A challenge to evolutionary theory?” from the book Perspectives on an Evolving Creation: Grand Rapids, and it coincides with our Question, "Does the Cambrian Explosion pose a challenge to evolution?". A pdf version of Miller's full paper can be found here.
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
“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.”
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.