It is very clear that most people who have posted to the blog site are quite uncomfortable with any proposed change to the concept of one big happy Anolis. What shines through to me in the posts is how deeply emotional the thought of this change is for many of us. I think I understand this emotion and hope to try to persuade you to let go of it by presenting this short story. I did my dissertation on Norops humilis in Costa Rica. The emotional side of me likes to think that, when this scientific name is mentioned in the future, my name and my work will forever be associated with it. Because of that, when Gunther Köhler and Kirsten Nicholson (my very own former student!!!!) wrote a paper demonstrating that I had not performed a dissertation on N. humilis but instead had worked with N. quaggulus, I took the news quite badly. In fact, to this day I struggle with this news because I find it difficult to deal with an emotion that says my work will be lost to the scientific community because of this name change. Obviously, this is totally illogical. The scientific community has been quite resilient to such changes. Classic works on North American Natrix were not lost to careful scientists by a name change to Nerodia. Blair’s work on North American Bufo will continue to be found and cited by anyone working with evolution of Anaxyrus. In the case of my N. humilis work, the thing that has gotten me over the emotional hump is the exciting biology that becomes clear if N. humilis and N. quaggulus are distinct species. Jenn Deitloff, Kirsten Nicholson, and I have been looking for the contact zone between the species I studied at La Selva and the species in Costa Rica that I thought I was studying. We want to determine how two species can maintain separate evolutionary trajectories given that there is no obvious boundary to their dispersal and their dewlaps, at least to my eye, are virtually identical. Köhler’s work seems to indicate that anole biologists have vastly undercounted the real species richness within Norops (and probably the other genera) because some characters, like dewlap color, may operate on a much more subtle level than we have allowed ourselves to consider. If I could have forced the world to succumb to my emotions, I would have, and these anoles would still be one big happy species rather than the several smaller lineages that character data seem to indicate they are. I could cling to N. humilis by pointing to a node on the tree and argue that, because of taxonomic stability, this should continue to be that species so that my La Selva work would maintain its association with that taxon. But, I would miss out on the interesting biology that emerges from simply letting go of that concept.
I see similar advantages to breaking anoles into eight genera. My experiences have caused me to develop a completely different search image for anoles in the genus Dactyloa than I have for those in the genus Norops. In helping to generate the revised taxonomy, I think I learned something interesting about anole ecology, and that is that it may be shaped by an origin of the group in the crowns of canopy rainforest trees in South America followed by a series of biogeographic events that brought them down to the leaf litter. I don’t recall our notions of evolution of anole communities being framed in quite this way. The fossil record and the topology of the phylogenetic tree led us to that insight. Discussions among the authors of the revised classification, during which we forced ourselves to use eight generic names instead of one, helped us gain those insights. We encourage the use of our taxonomy because it helped us see things that we might not have seen and we are confident that this may happen to others. As foundational as Schoener’s studies of one- and two-species islands were (and are – this work certainly shaped my interests), we think it would have been improved had he been forced to recognize those anoles as belonging to the genera Dactyloa and Ctenonotus. We suspect he would have analyzed the sets of islands separately and might have generated discussion among ecologists about degrees of freedom in comparative studies a decade before that discussion actually emerged. We think the taxon-loop vs. character-displacement argument would have been refined had the Dactyloa islands been viewed separately from the Ctenonotus islands. The Dactyloa-islands likely would have been described as fitting most strongly with the taxon loop hypothesis (large ancestors forced to become small with the first small species being doomed to extinction by the next smallest species – or large colonists reaching these islands, leading to the same process) and the Ctenonotus islands likely would have been described as most strongly fitting the character displacement hypothesis (mid-sized ancestors with a niche focused toward the ground diverging to make room for the next mid-sized colonists). We think Losos’ analysis of evolution of ecomorphology of Puerto Rican anoles would have been improved had he been forced to use the genera Deiroptyx and Ctenonotus.
I think the real intent of this blog is expressed in Glor’s posts. In my opinion, he is clearly asking the community of anole systematists to band together as a unified voice against acceptance of the proposed new taxonomy. Obviously, the community of anole systematists has never been of one mind on this topic and I would hope that the community would recoil at the thought that we ever should be. The notion that the world recently came to accept a single large genus Anolis as the only viable concept can be rejected by the observation that some in the community of anole systematists continue to publish under names such as Dactyloa, Norops, and Ctenonotus (e.g. Savage’s book). Given what is happening with so many other large, cumbersome genera, I think it is inevitable that a revised classification of anoles will happen and those who are fighting so hard to prevent it will find their careers intact when they cross that inevitable threshold. Once there, I think they will wonder why they fought so hard against change.
KIRSTEN E. NICHOLSON, BRIAN I. CROTHER, CRAIG GUYER & JAY M. SAVAGE (2012). It is time for a new classification of anoles (Squamata: Dactyloidae) Zootaxa, 3477, 1-108
- In Support Of The New Taxonomy - October 3, 2012
Rich Glor
Thanks for joining the discussion and for your insights on this topic. You’ve made some excellent points about the potential value of a revised classification. I also agree with your suggestion that emotional attachment to particular names can contribute to retention of taxon names that we’re better of letting go of. I don’t think, however, that such an emotional attachment is the main reason that some have called for the retention of Anolis. Instead, most people seem to be arguing that the amount of confusion and labor that would result from changing so many binomials is not worth the gain in phylogenetic information content associated with a formal generic revision. This is not an emotional argument, it is a practical one.
I hope that you’ll continue to weigh in on our discussions here at Anole Annals. While AA’s editors certainly have their own opinions, we would like this blog to remain what it has been for the last two years – namely, an open forum where anyone interested in anole research can post their discoveries and opinions.
Barnaby
” most people seem to be arguing that the amount of confusion and labor that would result from changing so many binomials is not worth the gain in phylogenetic information content associated with a formal generic revision.”
That’s a very clear statement about folks’ differing opinions regarding the revision. It is a practical and very reasonable argument. But that’s not to say that emotion is not very much involved. Many of the statements on this blog against the revision obviously inflate the amount of resulting confusion and minimize the phylogenetic information gained. Also, there is a tendency to focus on the short-term inconveniences while ignoring the long-term benefits. I think that both of these are largely emotional. I think that some are worried that this exceptional radiation of lizards becomes somehow less exceptional when provisioned into different general. That is too threatening to some, hence the emotional responses.
Rich Glor
I’m sure people’s emotions will inevitably get involved to one degree or the other, but I’m not sure this is a productive direction for the discussion. I believe that proponents of the new classification are arguing for their proposed revisions for scientific reason and I hope they’re willing to believe that I’m doing the same.
Barnaby
I don’t intend to discount your reasons for keeping the status quo. Your posts have been extremely useful and informative. There are certainly quite valid arguments for keeping Anolis sensu lato.
Rich Glor
If we can agree to set emotion aside, and the inclination to intuit the emotions of others in particular, I think we can have a productive conversation about some of the excellent points in your previous comment. Focusing on exactly how and why confusion might ensue from a new classification, or how phylogenetic enrichment might result (as Craig Guyer does in his post), seems like a great route to an enriching discussion. I hope we see more thoughtful posts and comments on this topic, and from a variety of different perspectives.
IanWang
I don’t mean for this comment, on its own, to be in support of one side or the other of this argument, but I feel compelled to point out something that might potentially be confusing to some readers of this blog. I think an important point is that this post begins with a story about the discovery that A./N. humilis and A./N. quaggulus are distinct species, but the decision to “lump” or “split” species is not analogous to “lumping” or “splitting” genera. We have defined species concepts to determine whether something constitutes a species (although there is some variation in different species concepts, these at least agree that there are certain criteria which delimit species); however, we have no clearly defined set of criteria for how these species should be included in genera (except that a genus should be monophyletic).
As an additional note, I’m not sure if the concluding paragraph was intended in this way, but it seems to suggest that the Anolis community doesn’t need to adopt a single taxonomy. This strikes me as a somewhat unusual suggestion, because one of the primary reasons for having scientific names is to provide a single identifier for a species, as opposed to the many potential common names it could have. Surely, taxonomy will have very little value to anyone if everyone chooses to use different names.
David Steen
Regarding your additional note, I think the subtext of the relevant sentences is that science advances through testing hypotheses, critical thinking, and debate. In absence of the one true answer, a unified voice should be avoided. Otherwise, the field (any field, for that matter) becomes stagnant.
Martha Munoz
When a debate revolves around where an aribitrary taxonomic break point in a flawed Linnean system should lie, and when the same points and counterpoints be stated and restated, then to me it has become stagnant. There are more interesting questions to be asked from Nicholson’s phylogeny than whether or not we should create new genera, especially when it would thoroughly disrupt an active and productive field.
Rich Glor
I agree that hypothesis testing, critical thinking, and debate are all vital to scientific advancement. I think its hard to argue, however, that anole research has suffered from taxonomically inflicted stagnation.
Yoel Stuart
Seconded. There are testable hypotheses for what a species is (at least if you’re a proponent of the Biological Species Concept). There are no such testable hypotheses for genera.
Thomas Sanger
I have previously voiced my concerns in detail and Rich’s comment summarizes them nicely. However, for the sake of trying to better understand your argument I hope that you can clarify your second paragraph for me. It seems like your argument distills down to the notion that use of Linnean ranks – in this case genera – can potentially guide our study of evolutionary patterns and processes. But to me it also seems that the critical element of your argument is the use of the phylogeny NOT the classification scheme that the taxa are arranged under. Now I may be way off here but while comparative biology once relied on Linnean ranks to formulate hypotheses about evolutionary processes we now rely more heavily on phylogenies to develop and test hypotheses. A robust anole phylogeny and modern comparative methods also didn’t exist early in Schoener’s research career or, presumably, during your dissertation so I don’t know if this is a fair comparison to make with modern research strategies. So if I understand your arguments correctly I think that I am still most comfortable formally naming clades while leaving “genera” out of the discussion as I do not see the benefits outweighing the risks.
Martha Munoz
It is interesting that you say you have developed a “search image for anoles in the genus Dactyloa” because I work on the cybotoids, a group that under Nicholson’s classification would be in its own genus. Perhaps Central American anolologists see different search images, but that necessitates creating breaks among Caribbean anoles that simply do not exist. The cybotoids are extraordinarily similar to A. distichus, also from Hispaniola, which the new classification puts in another genus. My field students typically require a few days to be able to tell them apart, and even after that they can be difficult to distinguish at a distance, not to mention further confusion with juveniles and females. The cybotoids are also really similar to other trunk-grounds from around the Caribbean. To justify a genus for clades Nicholson et al. consider morphologically divergent (ie: South American clades), it seems you are forced to overemphasize differences in clades that are morphologically quite similar because of monophyly.
Rosario Castañeda
The statement “My experiences have caused me to develop a completely different search image for anoles in the genus Dactyloa than I have for those in the genus Norops” might be a bit misleading. Although different search images for members of the Dactyloa versus Norops clades may be effective in Costa Rica, where Dr. Guyer seems to have had most of his experiences with anoles, they would not be very effective in South America. In Costa Rica, the Dactyloa clade is represented primarily by giant anoles (SVL > 100 mm) of the latifrons series (one other medium-sized species in this group, Anolis chocorum, barely makes it into southeastern Costa Rica). These large usually green anoles are readily distinguished from most Costa Rican members of the Norops clade, most of which are much smaller and usually brownish lizards. However, even in Costa Rica, there is a relatively large and bright green member of the Norops clade, Anolis biporcatus, that was formerly thought (i.e., prior to Etheridge’s work) to be closely related to South American species now placed in the Dactyloa clade (e.g., Stuart, 1955). Moreover, in South America, there are various species in the Dactyloa clade, for example, A. festae and A. peraccae, that are not particularly large (SVL < 60 mm), are greyish or brownish in coloration, and thus bear a stronger superficial resemblance (the basis of a search image) to various members of the Norops clade than to the giant Costa Rican anoles of the latifrons series.
Stuart, L. C., 1955. A brief review of the Guatemalan lizards of the genus Anolis. Misc. Publ. Mus. Zool. Univ. Mich. 91:1–31.
(Co-written with Kevin de Queiroz)
Levi Gray
Producing a drawn-out ad hominem suggestion that emotion is one of the major factors keeping workers opposed to the Nicholson et al. taxonomy is quite frustrating to those that wish to have a rational discussion on the topic. The arguments against accepting the changes have come from many angles, none of which are driven by emotion.
If you want to know why so many oppose the new taxonomy, look to the many legitimate arguments put forth by those on this blog. Anyone that wishes to ignore those arguments due to some perceived “emotional attachment” to the genus Anolis or opinions of past advisors needs to take a moment to consider what has actually been said, and by whom. These criticisms come from a variety of different fields and academic backgrounds. They address the methods, practicality, and taxonomic philosophy of Nicholson et al. It would be appreciated if those issues were addressed directly, instead of something that we can all agree does not belong in these arguments (emotion).
Kirsten Nicholson
Levi and others, in reference to the “emotion”: I think it has more to do with the way some of the comments and posts are written which sounds emotional to many of we readers. Several people have commented to me and not posted on here regarding the “tone” of the posts. Its clear that folks on both sides of the argument are passionate about what they believe in. And I think that’s part of the problem: trying to make an objective, scientific classification system that all can follow and understand and has practical application for us all, rather than a system that requires users to “believe” it should be one way or another.
Shea Lambert
I think part of the problem is that there is no truly “objective” way to assign the Linnean rank of genus — whether we like it or not, assigning new genera is an activity wrought with subjectivity, and it could have been done for Anolis in a thousand other ways (or not done at all). In my view, nomenclature is not science; we cannot test whether the eight new genera are actually genera or not. This debate is necessarily subjective and boils down to “which option will result in better science?” I’m not convinced that it is valuable or fair to imagine what effects these new genera would have had if applied to work from 20+ years ago, and think that we should focus on how these different options for nomenclature will affect research moving forward.
Shea Lambert
I’d just like to add that, because these decision is entirely subjective (outside of the requirement that genera be monophyletic), I don’t think that arguments regarding consensus and the scientific process (e.g., last paragraph of Craig Guyer’s post, comment by David Steen, tweets from kwren88) really apply here. We can all have different opinions on which particular bifurcations on the tree are important, and that’s good for science, but I fail to see how disagreement on nomenclature itself is going to benefit the anole research community.
Shea Lambert
I realize now that I’ve misused the word “nomenclature” in my above comments and that “Linnean classification” should probably go in place of it.
Yoel Stuart
Craig, you and I are friends. So I hope that you know that I’m asking the following question sincerely. If you take your arguments to the logical conclusion and the splitting continues according to finer and finer distinctions until all anole species are in monotypic genera and your graduate-school-near-and-dear genus Norops is reduced to containing one species, Norops humilis even, would you be okay with that?
If yes, then you truly contain no emotion about this and I suspect that you and I will never see eye to eye on this. We’ll agree to disagree.
If no, then when would you stop the splitting? 8 genera, for now. There are 22 well supported clades that could also get generic names. Why not those? If you eventually get to those, why not more? At some point you’d make a subjective decision to stop. And a subjective decision is an emotional one.