We’ve already had lots of discussion about Nicholson et al.’s (2012) recent proposal that Anolis be fragmented into eight genera. Throughout the course of this discussion, several posts and comments have suggested that anole biologists might be compelled to implement Nicholson et al.’s proposed generic revision by the International Committee on Zoological Nomenclature (ICZN) and its rules for nomenclature (a.k.a. the ICZN* or The Code) (see comments on recent posts by Losos and Sanger).
Although I must admit at the outset that I am not an authority on The Code or its implementation, I will argue below that the belief that the code compels anole biologists to accept Nicholson et al.’s proposed taxonomic revision is completely false. The ICZN has neither the authority, nor the interest in, policing taxonomic practice and will have no role in determining whether Nicholson et al.’s (2012) new generic classification is accepted or rejected by the community of researchers who study anoles. I believe that the reasons for this are fairly straightforward and uncontroversial, but they do require us to think a little about our taxonomic philosophy and the difference between taxonomy and nomenclature.
Let’s start with some basics for the non-systematists. According to the ICZN, the goal of taxonomy is “the identification and interpretation of natural groups of organisms (i.e., taxa) based on characters (such as morphology, genetics, behaviour, ecology).” One piece of good news for anole biology is that everyone involved in debate over Nicholson et al.’s new classification shares the same fundamental taxonomic philosophy – namely, that taxa should be diagnosed using phylogenetic trees and should correspond with monophyletic groups. We may debate whether certain taxa are supported as monophyletic by the available data, but we all agree that recognition of monophyletic groups is a primary objective of any taxonomic scheme for anoles.
More good news: The Code has no interest in getting involved with taxonomic decisions. I realize that the The Code can be really boring to read, but you don’t have to read more than the first two paragraphs of the introduction to get this message (in a few cases I’ve added my own emphasis by bolding text):
“The 4th edition of the International Code of Zoological Nomenclature … has one fundamental aim, which is to provide the maximum universality and continuity in the scientific names of animals compatible with the freedom of scientists to classify animals according to taxonomic judgments. The Code consists of Articles … [that] are designed to enable zoologists to arrive at names for taxa that are correct under particular taxonomic circumstances. The use of the Code enables a zoologist to determine the valid name for a taxon to which an animal belongs … There are certain underlying principles upon which the Code is based. These are as follows: (1) The Code refrains from infringing upon taxonomic judgment, which must not be made subject to regulation or restraint…”
Rather than concerning itself with taxonomy, which inevitably involves subjective decisions made by systematists that specialize on particular groups of organisms, The Code focuses exclusively on nomenclature, or “the system of scientific names for taxa (such as species, genera, or families) and the rules and conventions for the formation, treatment, and use of those names.” The Code, therefore, merely provides “a set of rules for the naming of taxa that follows an internationally agreed, quasi-legal procedure.”
With this background, we can return to a consideration of Nicholson et al.’s classification and the role that The Code may have in its implementation. Nicholson et al. argue that in order to appreciate and study the phylogenetic diversity of anoles we must formally recognize the taxon that includes all anoles not as a single genus, but rather as a number of related genera. Although determining whether to proceed with the traditional classification involving a single genus or the Nicholson et al. classification that recognizes eight genera might seem to be a distinction between two alternative systems of nomenclature whose outcome is dictated by the The Code, this is not the case. Instead, both alternatives are perfectly compatible with The Code, and the decision about which classification to adopt moving forward is a subjective taxonomic decision that must be made by the community of biologists who study anoles.
All The Code says is that if we anole biologists want to recognize the taxa that Nicholson et al. have diagnosed as genera, we must use the names they have resurrected from the historical literature and applied to these taxa. If I wrote a paper tomorrow that gave a new generic epithet to the same taxon that Nicholson et al. have named Ctenonotus, this new name would be rejected under the rules of priority outlined in The Code. However, The Code respects the right of anole biologists to make the subjective taxonomic decision about whether we want to recognize the taxa diagnosed by Nicholson et al. as genera, or instead recognized them informally as series or species groups, as anole biologists have done for decades. Recall from our earlier passage from The Code that its rules for nomenclature only apply “under particular taxonomic circumstances.”
My fellow anole biologists, we have a taxonomic decision to make and the ICZN is not going to make it for us. It seems that the worst outcome would be fragmentation of the community of anole biologists, with some researchers using the traditional approach and others applying Nicholson et al.’s revised generic classification. More readings and notes are after the fold.
For those who are interested in learning more about the taxonomic issues raised in this post, I encourage you to check out a few other recent papers on this issue:
de Queiroz, K., and M. J. Donoghue. 2011. Phylogenetic nomenclature, three-taxon statements, and unnecessary name changes. Systematic Biology 60:887-892 [doi link].
Pauly, G. B., D. M. Hillis, and D. C. Cannatella. 2009. Taxonomic freedom and the role of official lists of species names. Herpetologica 65:115–128 [doi link].
Hillis, D. M. 2007. Constraints in naming parts of the Tree of Life. Molecular Phylogenetics and Evolution 42:331-338 [doi link].
* Yes, an organization whose only goal is to encourage clarity in the application of names uses the same acronym for two different things; ICZN stands for both the organization known as the International Commission on Zoological Nomenclature and the set of rules known as the International Code of Zoological Nomenclature.
- JMIH 2014: Early Records of Fossil Anolis from the Oligocene and Miocene of Florida, USA - August 13, 2014
- JMIH 2014: Relative Contribution of Genetic and Ecological Factors to Morphological Differentiation in Island Populations of Anolis sagrei - August 7, 2014
- JMIH 2014: The Ultrastructure of Spermatid Development within the Anole, Anolis sagrei - August 5, 2014
Martha Munoz
While I totally agree that the “worst outcome would be fragmentation of the community of anole biologists, with some researchers using the traditional approach and others applying Nicholson et al.’s revised generic classification,” this has already been happening. Many times in Central America I have had heard taxa referred to exclusively as Norops or Dactyloa, without the qualifier that they are not genera, but rather clades within Anolis. I am familiar with the group and its systematics, so this doesn’t affect me that much, but what about the plethora of biologists who go down there, as well as ecotourists? What about people reading the smattering of papers that utilize these terms and cannot piece together the history of the field? We’ve obviously touched on these points before.
Obviously, there is a very specific school of thought there, founded and expounded by the very people writing this paper. From a scientific perspective, how do we achieve consensus on this issue? And even if we can, through majority rule and cogent argumentation, dispel the idea that 8 genera are the way to go, how do we get everyone on board? Other than publishing papers and using public media such as this blog, is there a forum for the field to make a decision on this issue?
Rich Glor
Thanks for your feedback Martha. The issues you raised are the reasons we’re hosting a week-long discussion of the Nicholson et al. monograph! I hope that people will be able to hash out the plusses and minuses associated with each alternative and leave the week with a clear sense about the best path going forward. Tomorrow (Tuesday), we’ll have some new perspectives on the plusses and minuses of Nicholson et al.’s proposed taxonomic revisions.
James Jackson
“Obviously, there is a very specific school of thought there, founded and expounded by the very people writing this paper.”
What are the premises of this school of thought? As I pointed out in my comment of September 23, 2012 at 6:49 am on an earlier post, Dr. Crother co-authored an opinion paper which argued that elevation of a set of taxa from a lower to a higher categorical level can reduce the information content of the classification. Why is that premise valid for the classification of North American snakes but not for anoline lizards? What am I failing to understand here?
A semi-objective criterion for allocation of monophyletic taxa to categories could be to attempt to keep morphological/ecological disparity similar across a given category. For example, if Sceloporus were taken as “the disparity standard” for iguanids, then the study by Warheit et al. could perhaps argue for recognizing multiple anoline genera. Whether eight genera would be supported under this criterion would require assessing disparity within those proposed genera and comparing the results with Sceloprous disparity.
Martha Munoz
Hi James,
I apologize for any potential confusion. By school of thought I was actually referring specifically to Jay Savage and his students. The “premise” for this school of thought is to rewrite Anolis taxonomy around their conception of Norops as a valid genus. Please see Rich’s post today about the history of Norops and Anolis.
To me Savage and colleagues have a long-standing agenda to elevate Norops to a genus. If that involves separating alpha and beta anoles into separate genera, so be it. If, in light of evidence that Norops renders Anolis paraphyletic, and so we must now build 7 other genera around it, so be it. Whatever it takes.
The “semi-objective criterion for allocation of monophyletic taxa to categories” is more simply accomplished by naming clades within Anolis rather than new genera, in my opinion.
James Jackson
Your hypothesis that there has been a long-standing agenda to elevate Norops to generic rank is, of course, consistent with Crother’s not applying the “information content criterion” even-handedly both to North American snake classication and to anoline lizard classification.
Regarding classication, perhaps biology needs parallel systems: Linnean for generalized communication with the non-evolutionist public (which would also include some biologists), and PhyloCodean for communication among students of the evolutionary process. For example, the kind of question that the latter should be fascinated by is why, counting back six nodes from Enyalius pictus, you cross a treshhold to a more greatly different morphology (Anisolepis/Urostrophus) than if you count back six nodes from, say, Anolis carolinensis.
PhyloCode nomenclature would probably be superior to Linnean for discussing answers to my hypothetical ecomorphological/ecoevolutionary question.
Liam Revell
Nice post, Rich.