As we finish up combining the data, it’s time to start thinking about the specific analyses that we’re going to do. What are the specific questions we’re asking? What are the techniques that we need to address the questions? Some excellent discussions between ODPers have been happening in one of the recent posts, and I was hoping to continue that here. In particular, I wanted to refocus the discussion on the project’s essential questions, and consider the types of analyses that we can use to answer each question. I’m just thinking out loud here (this is open notebook science, after all), and invite suggestions and discussion in the comments section. In particular, I’m referencing the “big questions” outlined in one of our first posts.
Why did ornithischians evolve quadrupedality multiple times?
I think this one is going to have to simply rely upon our interpretation of the data. After all, we can perhaps answer “how,” but the “why” can’t really be answered in this setting. So, it’s something to consider in the “discussion” section of the paper. But, see the next question. . .
Was the evolution of quadrupedality consistently associated with an increase in body size?
Here, we’re basically looking at evolutionary trends. In other words, can we detect a trend in body size within various ornithischian lineages? The more I think about this, the less I’m convinced we can directly answer the question (if you disagree, and have a solution, pipe up in the comments, please). One problem is the difficulty in knowing whether or not certain taxa were truly quadrupedal. So, where do you make the cut-off for quadrupedal vs. bipedal vs. both? In many cases we just don’t know. There’s a danger in circular reasoning, too (the limb bones look like it’s quadrupedal, so we call it quadrupedal, and then use it as an example of a quadrupedal taxon for analysis of limb bones).
But, I think we can detect trends across Ornithischia as a whole, and within specific lineages. For instance, is there a trend for increasing body size across Ornithiscians? Is there a trend for increasing body size within Ornithopoda? Ceratopsia? Thyreophora? In fact, Matt Carrano found a consistent and statistically significant increase in body size within ornithischians (and indeed, within most dinosaurs) when considering femoral measurements (go here to download a free PDF of Carrano, 2006). So, that makes this question a little less interesting (and indeed, less publishable, because it’s already been done). Do you think we should move it to the back burner? Or should we spin it in another way? Thoughts are welcome.
Did different groups of quadrupedal ornithischians arrive at this body form in similar ways, or did they have different strategies?
Here (as far as I know) is a genuinely novel question, and I think it’s the core of the ODP’s current phase. What we’re really saying (I think) is this: We know that thyreophorans, hadrosaurs, and ceratopsids independently evolved quadrupedal locomotion. Did each group have similar limb proportions, or were they different? I think this is where we’ll want to look at principal components analysis, at least as a starting point for data visualization. And, we’ll have to do that within a phylogenetic context. A recent paper by Liam Revell (2009) addressed how to do this (thanks to ODPer Randy Irmis for bringing up this paper; you can download it for free here – it’s well worth a read).
A second way to look at this question is to look for trends in certain structures – for instance, do the metacarpals tend to get elongated in each group (relative to the rest of the arm) as different clades became quadrupedal? Here, we might use a simple non-parametric correlation of the ratio with patristic distances (see the Carrano paper, again, and references therein, for a brief introduction to this method), to investigate that question within different lineages. Basically, patristic distance estimates the distance of a particular species from the base of the tree (by the number of branching points leading up to it). A taxon that split off early in a group’s evolution would have a low patristic distance, and vice versa for one that split off late in a group’s evolution. So, we might look at the correlation of metacarpal:arm length ratio to patristic distance for thyreophorans, hadrosaurs, and ceratopsians.
I think I’ll end here for now! Please add thoughts, suggestions, corrections, and anything else you think relevant in the comments. Next time, I’ll move on to the final issue, quantifying morphological disparity in ornithischian evolution.
Carrano, M. T. 2006. Body-size evolution in the Dinosauria. In M. T. Carrano , R. W. Blob, T. J. Gaudin & J. R. Wible (eds.), Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles. University of Chicago Press, Chicago:225-268. Freely available here.
Revell, L. J. 2009. Size-correction and principal components for interspecific comparative studies. Evolution 63: 3258-3268. Freely available here.