Transforming Our Thinking about Transitional Forms

The concept of a “missing link” is an “archaic expression” (Padian and Angielczyk 2007) tracing back to the Great Chain of Being, a view of the physical and metaphysical world as an unbroken chain. It was later temporalized by the evolutionary thought of the eighteenth and nineteenth century to the idea of evolution as a progressive climb up a ladder (Ruse 1997). These views of evolution create the false expectation that there should be fossil evidence showing “a complete chain of life from simple to complex” (Vardiman 2003:328). Creationists rely on such views to support their arguments against macroevolution, in particular by pointing out the “conspicuous” absence of “large numbers of intermediate fossil organisms” (Changing lives 2003), using what is still unknown to question whether evolution has occurred. I will deal with the misconception of evolution as a ladder-like progression shortly, but should the fossil record be expected to reveal all species that have ever lived? Clearly not. Knowledge of the fossil record will never be comprehensive (Darwin 1859; Padian and Angielczyk 2007; Prothero 2007). First, there is too much of the Earth to explore, and paleontologists have to be content with samples. Second, given our knowledge of geology, we understand that not all organisms will be fossilized and that there will be systematic biases in what organisms are fossilized (Kidwell and Holland 2002). Therefore, any statements rest on a fallible, if informed, assessment of the necessarily incomplete evidence.

Even if one does not expect that the whole Great Chain of Being should be found in the fossil record, there is yet another outmoded view of evolution driving creationist antievolution propaganda. Expectations of “intermediate” forms reflect neither Darwin’s original thoughts nor current thinking and practices in evolutionary biology and paleontology. Darwin (1859) was very clear on this. The ancestor of two living forms is unlikely to be found alive because it would have been outcompeted in most cases by newly adapting forms, and an extinct ancestor of two living forms would not be expected to look intermediate between them. Today, evolutionary biologists and paleontologists do not focus on finding “intermediates” but rather on reconstructing evolutionary relationships and history using shared derived characters or synapomorphies. Willi Hennig revolutionized systematics in the 1960 s with the introduction of cladistics, which ushered in a new method of phylogenetic analysis and a new approach to systematics. Instead of relying on a Linnaean system of classification, cladistics placed the focus on evolutionary history, specifically identifying features as ancestral (general) or derived (evolved after the lineage split from the ancestor). If a shared derived character or synapomorphy is found in two or more related organisms, it is inferred to have been present in their common ancestor, irrespective of whether or not there is a fossil record for that ancestor (Hennig 1966). Rather than trying to find the actual fossil corresponding to the “missing link” between lobe-fins and tetrapods, paleontologists instead look for fossils with characters or features important for an adaptive transition from life in an aquatic environment to life on land and that are shared as the result of common ancestry.

David Attenborough recently declared in a program about Darwin that “you’d have to be extraordinarily blinkered if you didn’t stub your toe against the theory of evolution very early on in your life”, particularly if as a child you collected fossils (Charles Darwin and the tree of life 2009). The data to substantiate many of the major transitions in evolution have been amassed. Lines of evidence include cladograms from independent datasets (i.e., fossils, morphology, and molecules), stratigraphy and radiometric dating to establish events in time, and paleoecology to interpret ancient environments. Embryology, behavior, histology, geochemistry, functional morphology, physiology, and biochemistry all add to the array of independent lines of evidence used to test evolutionary hypotheses.

Padian (2008) suggests depicting these various lines of evidence, particularly the ones that are important in major evolutionary transitions, in a single diagram, called an “evogram” (Fig. 3), to help students to understand that evolutionary research is conducted in an integrative way and in a way that relies on testing independent lines of evidence against each other to demonstrate concordance. Evograms have a cladogram as the backbone of the diagram, generated using an entirely separate base of information. Because a cladogram is basic to understanding evolutionary relationships, other types of noncladogram evolutionary diagrams will only confuse students (Catley and Novick 2008). In Fig. 3, fossil evidence of limb structure is depicted in one row. Structures of the fossils, showing corresponding bones in corresponding colors, are interpreted in the next row. Important clade names and important synapomorphies can also be added to the nodes of the cladogram (Padian 2008). Evograms integrate the various independent lines of evidence, providing a more united picture of evolution.

The challenge is that teachers are generally neither trained nor equipped to teach this way. Indeed, tree-thinking has only recently trickled down to high school biology (Goldsmith 2003; Catley 2006; Baum and Offner 2008). Most high school and many college introductory biology texts are woefully lacking in their presentation of comprehensive figures integrating phylogenetic analyses with hypotheses about evolutionary process. In fact, it appears that many of the diagrams may actually be misleading (Catley and Novick 2008). In a recent analysis of evolutionary diagrams in biology textbooks, Catley and Novick (2008) found that even though 72% of the diagrams in their survey of 31 textbooks were cladograms, there were striking differences across grade levels. Middle school texts had the fewest cladograms, with almost twice as many noncladograms. High school texts included more cladograms, but the ratio of cladograms to noncladograms was approximately equal. Such high proportions of noncladograms are disturbing if these support misconceptions about evolutionary processes (Catley and Novick 2008).

I urge teachers to embrace cladistics and tree-thinking as a way of examining the natural world and encourage students to make such thinking a habit. For a first step, the following websites offer basic exercises on cladistics and tree-thinking: Evolution and the Nature of Science Institutes (http://www.indiana.edu/~ensiweb); Understanding Evolution (http://evolution.berkeley.edu); What did T. rex taste like? (http://www.ucmp.berkeley.edu/education/explorations/tours/Trex). Also recommended are ways of teaching about evolution consistent with modern practice in systematics: using “ancestral” and “derived” instead of “primitive” and “advanced” when referring to organisms and characters, which promotes thinking about evolution as a tree rather than a ladder (and also follows, the good example of Darwin, who admonished himself, “Never use the words higher or lower”); including exercises that define clades by synapomorphies and map such characters onto phylogenies (see Staub et al. 2006 and ENSI: Making Cladograms); and clarifying that a cladogram or phylogenetic tree is a hypothesis, given the data, and that a node on a cladogram is a hypothetical ancestor, not an actual ancestor for which we should expect to find a fossil representative. To attain scientific literacy, students need to learn that evolution is a branching process, that paleontologists are not searching for missing links but for transitional features, and that evolutionary research is conducted in a way that enables scientists to use independent lines of evidence to converge on robust conclusions about the history of life. Teachers need to learn these lessons in order to be able to impart them!