Year: 2012 Page 14 of 47

What’s In A Name?: Scientific Name Use For Anoles, By The Numbers

As should be evident from several recent Anole Annals posts and comments, Nicholson and colleagues published a paper last week proposing that “It is time for a new classification of anoles.” Among a number of arguments in favor of splitting up the genus Anolis, Nicholson et al. (2012) argue that use of a single genus name hinders scientific communication about these animals. This argument has generated a lot of discussion (e.g., a post by Sanger, and two different threads of comments found here and here), and I thought it might be useful to continue the discussion with a bit of information about the usage of anole names in the scientific literature.

In a comment on an earlier post, Duellman argued that a genus name does not simply exist to reflect systematic knowledge – it’s a (hopefully stable) handle that conveys information about identity to a very wide audience, from laypersons to college students to ecologists, conservationists, and systematists.  My impression has always been that this is especially true for Anolis – more so that for many other groups of organisms. For example, geckos are commonly known, even to scientists, by their common name “gecko,” and we find this term in paper titles and abstracts. I don’t think this is true for anoles – it seems to me that we more often simply call them “Anolis“.

To see if this is actually the case, I decided to pull some numbers from Web of Knowledge. I conducted a series of “Topic” searches for various taxonomic names, such as “Anole”, “Anolis“, “Gecko”, etc., and recorded the numbers of matching records for each search. Records include instances in which a term is found in the title, keywords, or abstract of any book or article recorded in the Web of Knowledge academic database. The numbers returned are reflective of my university’s library holdings (University of California), and will be different if conducted elsewhere; I also didn’t spend any time processing the results, but I don’t think that should qualitatively affect any results.

The Case For Splitting Up Anolis

ResearchBlogging.orgPrevious posts on AA are engendering a lot of discussion about the proposal to reclassify Anolis into eight genera. Because most of the comments are critical, we felt the positive side of the case should be presented explicitly to AA readers. What follows is a summary of the arguments in favor of dividing Anolis into eight genera, drawn primarily from Nicholson et al.’s paper.

The argument for splitting Anolis is straightforward and is laid out clearly in the paper (p.13): “The role of systematics is to advance our understanding of biological diversity in the natural world. Its practitioners are the guardians of the knowledge produced by past generations and responsible for the rational interpretation of new data and their implications. Within this framework, phylogenetic inference has consequences that we think bind its practitioners to produce a systematic classification of the studied organisms. Such a classification must be founded on the inferred evolutionary relationships and dictated by the canon of monophyly. Following the above precepts, in conjunction with our phylogenetic analyses, we recognize eight major evolutionary units (genera) and twenty-two subunits (species groups) of dactyloid lizards (Figs. 4–5). The current practice (following Poe, 2004) of treating all dactyloids as comprising a single genus underemphasizes the evolutionary diversity within the family (as currently recognized) and obfuscates major biological differences among clades. In addition, simply because of the large size of the family (nearly 400 valid species), the single genus concept can be a hindrance to scientific communication regarding evolutionary events and directions of future research.”

In other words, the authors argue that failing to recognize structure within the anole clade obscures knowledge of phylogenetic relationships. If we can identify such clades, we should give them generic status to promote dissemination of this knowledge. Todd Jackman, though somewhat neutral in his stance, concurred with the rationale in a comment yesterday (comment #2): “I would like anyone working on anoles to know these eight groups, and to be familiar with the 22 subclades as well — but how to best achieve better knowledge of the phylogeny of anoles is not straightforward. Using subgeneric or clade names is fine, if they get used and get used often. If only taxonomists and serious tree-making anole workers use the names for these clades, then the phylogenetic information hasn’t been conveyed.  Splitting up the genus…forces everyone to use more phylogenetically precise language.” Looked at another way, our best hypothesis of anole relationships reveals eight clades. By highlighting these clades with generic status, we explicitly put them forth as a hypothesis for future testing and potential falsification. The authors conclude that failing to do so stymies systematic progress (p.4): “Systematic progress in this regard has been delayed by an extremely conservative taxonomic approach to recognizing the diversity within the group and its extraordinarily ancient historical roots.”

In addition, a genus of 400 may be unwieldy. How can one easily distinguish anoles that are closely related from those that are more distant? Lumping them all in one genus might obscure information and thus obscure evolutionary patterns and lead to inefficient or even misguided choices in research design and interpretation.

Finally, retaining a large—and very old—genus Anolis runs counter to prevailing practice these days, which is to split rather finely, producing genera that are young in age and with relatively few species. As a result, Anolis is an outlier, being very old (100 million years plus, according to this paper). Some—we won’t name names—have been known to crow that Anolis is the most species-rich amniote genus, but that’s not very surprising if Anolis evolved tens of millions of years earlier than other genera. Many in the community feel that old genera should be split up, a view shared by AA reader Barnaby (currently comment #5 in the string).

For these reasons, Nicholson et al. suggest dividing Anolis into eight genera.

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

Anolis: Should It Stay Or Should It Go?

ResearchBlogging.orgNicholson et al. recently undertook the bold mission of revising the taxonomy of our well-loved lizard genus, Anolis, based on the phylogenetic relationships among its many species. Not surprisingly this has struck a nerve with much of the anole community spawning a range of reactions immediately following its publication, some applauding their efforts but many expressing their concerns about the proposed change. If one of the author’s objectives was the generate discussion on this topic its clear that they have succeeded.

The Nicholson team should first be commended for their efforts to synthesize the historical literature on anole taxonomy, encompassing “387 recognized species and 112 additional nominal subspecies” with some reports dating as far back as the mid-1600s. This survey will likely serve as a benchmark for later systematic evaluations of this genus. However, the implications for their proposed revision extend well beyond the nuances of taxonomic rule or the analytical methods used to build phylogeny*. The issues arising extend into other biological disciplines and potentially undermine the rich intellectual history of anoles.

I, like many others, am a consumer of taxonomy and systematics. These are critical to the comparative analyses I perform and in communicating my findings to others in the anole community, herpetologists more generally, and other biologists more broadly still. Anolis has been a model for comparative biology for decades but is gaining increased attention by genomicists, neuroendocrinologists, and developmental biologists. Just this year, in fact, the anole community developed a system with which to share comparative molecular resources. Deconstructing Anolis into eight distinct genera could drive an intellectual wedge between the previously published literature and future studies, potentially derailing the continuity of information that is critical for academic advancement. This change could lead to unforeseen consequences that damage the broad utility of Anolis among biological disciplines that depend on the stability of anole nomenclature.

Nicholson et al. state, “the role of systematics is to advance our understanding of biological diversity.” While I agree with this statement in principle I feel that it is also important to ask if the benefits of revising this diverse taxon outweigh the risks I outlined above. The glaring disconnect between phylogenetic systematics and Linnean ranks is discussed at great length elsewhere and will be strategically avoided here. It is worth asking, however, whether the addition of new genera (specifically genera, not simply clade names) add anything new to our biological understanding of this group. Ultimately, can we more accurately communicate our findings using the revised nomenclature? While Nicholson et al. use monophyletic clades to distinguish the proposed genera  – a well respected practice – the precise breaks are biologically arbitrary. In my opinion the suggested genera do not offer greater clarity to the natural history of this clade as they do not partition Anolis based on distinct biogeographic groups, groups with distinct ecologies, or groups with distinct, readily recognizable morphological features. In this proposed taxonomic scheme the ecological and morphological convergence of Anolis ecomorphs** that is widely discussed and cited throughout ecological and evolutionary literature becomes a confusing hodgepodge of convergent lineages from different genera. In my opinion it is overwhelmingly clear that the benefits of re-classifying Anolis lizards do not outweigh the ensuing upheaval of our research community.

At face value it appears that the overall motivation for revising Anolis is its diversity, as it is undoubtedly one of the most diverse tetrapod genera. However Anolis pales in comparison to many invertebrate genera. The beetle genus Agrilus (jewel beetles) has an estimated 2886 species! Drosophila – the genus that possesses the genetic and developmental powerhouse D. melanogaster – contains approximately 2000 species***. It is clear that large, active research communities can readily work with diverse genera without problematic communication of their results. The sole argument of diversity is not strong justification for revising Anolis.

Perhaps some day taxonomy will abandon the binomial naming scheme derived from the Linnean classification hierarchy in favor of a more accurate system based solely on phylogenetic systematics. However, for practical purposes, we are simply not there yet. Anolis serves as a great example of where premature taxonomic revision could have far reaching consequences that can send biological research in multiple disciplines into severe turmoil.

Comments and discussion on the ideas I have shared above are welcomed and encouraged!

* This is not the say that critical evaluation of phylogenetic methods are not essential to the evaluation of taxonomic hypotheses. I will save evaluation of the Nicholson et al. analyses to those with greater experience working this these methods and those with an intimate knowledge of the proposed species groups.

** Beyond their proposed taxonomic revision the Nicholson team also reject the Anolis ecomorph concept. This idea will no doubt attract additional attention from the community.  Stay tuned to Anole Annals for more on this issue.

*** A similar discussion to ours recently took place in the Drosophila community and many of these same concerns were expressed. O’Grady and Markow 2009 state that “such radical taxonomic revision is not advisable…as the literature and traditions are
so well established that any such formal reassessment would not be worth the confusion engendered.” After review and comments from the community the ICZN voted that taxonomic revision of Drosophila was “premature” and wisely left this diverse genus intact.
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

The Proposal To Split Anolis Into Eight Genera: Time To Discuss

ResearchBlogging.orgWe’ve had a week now to let the proposed reclassification of Anolis sink it, so it’s time to start discussing it. A revolutionary new view of the scientific review process suggests that in the future, all papers will be published open access online (as this one is–thanks Nicholson et al.), the journal in which it appears (if any) will not matter, and peer review and evaluation will be conducted post-publication on internet discussion sites. Realistic? Who knows, but why not give it a try?

The paper by Nicholson et al. is undoubtedly the most important paper on anoles to be published in the last several years. Not only does it propose to split Anolis into eight genera, but it also presents provocative findings about the ecological evolution of anoles (including throwing out the ecomorph concept), anole biogeography, and the dating of evolutionary events in anole history.

Anole Annals’ goal is to be the meeting place for discussion of all things Anolis, so let’s take this post-publication review and commentary idea out for a spin. Anole Annals invites members of the anole community to post their thoughts on any aspect of the Nicholson et al. paper. We hope to get a conversation going on the merits of splitting the genus, as well as the other issues raised in the paper. In fact, this has already begun, as evidenced by the comments by Mssrs. Crother, Hillis and Duellman, among others.

To get the ball rolling, here’s a short précis of the paper:

1. Phylogenetic analysis based on previously published data of all sorts (genetic, morphological, karyological), with a smidgeon of new molecular data, reveals a phylogeny with eight strongly supported clades in a Bayesian analysis. These clades are recognized as distinct genera.

2. The ecomorph concept does not apply to mainland anoles because species similar in habitat use are not similar in morphology. Hence the term “ecomode” is coined for species similar in habitat use. Phylogenetic analysis of ecomode evolution on the phylogeny suggests that the crown-giant ecomode is ancestral for Anolis. The ecomorph concept is argued to not work for Greater Antillean anoles and should be discarded.

3. Biogeography is reconstructed on the phylogeny. Using the phylogeny, the authors argue that the eight clades differentiated about the time that the proto-antillean islands were passing between what is now North and South America. The Norops clade differentiated on several of these blocks (both island and mainland), explaining why Norops is nested within Caribbean non-Norops taxa without requiring the island-to-mainland colonization of Norops proposed by a number of previous papers.

4. Molecular clock dating reveals that anoles are surprisingly ancient, originating in South America approximately 130 million years ago.

Nicholson, K. E., B. I. Crother, C. Guyer, J. M. Savage (2012). It is time for a new classification of anoles (Squamata: Dactyloidae) Zootaxa, 3477, 1-108

Anolis Aequatorialis Information

Photo by Jonathan Losos

I’ve just learned of a website called Atyourservices. I don’t understand the point of this site, which seems to have a disparate range of topics, but one of the posts today is a nice compilation of sources of photos and information on Anolis aequatorialis.

The Amazing Social Life Of The Green Iguana

From http://blogs.scientificamerican.com/tetrapod-zoology/2012/09/17/amazing-social-life-of-green-iguana/

Here at Anole Annals, we occasionally digress to post on interesting topics in anole relatives. In that vein, I wish to call attention to a fascinating summary of the social complexity of Anolis‘s big green cousin, Iguana Iguana. Tetrapod Zoologya fascinating source of information on all thing Tetrapodan, has a very interesting article which I highly recommend.

Explain What’s Going On Here

I’ll give one hint: it started something like this.

Anolis Gemmosus

Those guys at Tropical Herping have done it again. Check out their new post with information and lovely photos of Anolis gemmosus, truly a gem of a lizard.

Jumping Without The Tail Is Bad For An Anole, And It Might Not Get Better

ResearchBlogging.orgAn interesting paper in 2009 showed us that jumping without a tail can be a disaster for green anoles. In that paper, the authors found that the bodies of tailless individuals often underwent extensive posterior rotations in the air, resulting in very awkward landings. Moreover, tail regeneration can take months to complete, which implies that losing stability in the air may not be a short term situation. So we wondered: can green anoles quickly improve in-air stability, or do they just have to wait until they have their tails back again? To address this question, we tested in a recent study whether tailless green anoles can improve in-air stability in five week’s time and whether gaining more jumping experience facilitates the improvement.

We found that there was extensive variation in how much an individual could improve within five week’s time. By the end of our study period, some individuals showed no sign of improvement,

httpv://youtu.be/aTfui2FlC9Q

whereas others did improve their in-air stability as time went by.

httpv://youtu.be/5s86p8KMWTE

Interestingly, the acquisition of more jumping experience did not seem to matter. Lizards with more jumping experience on average did not do better than those without. It appeared that the motor coordination capacity of an individual might be the most relevant factor for locomotor recovery in tailless green anoles. Our finding suggested that the cost of tail loss might be very different among individuals in natural populations. It would be very interesting to perform a manipulative field study to see whether individuals that are unable to improve in-air stability alter their habitat use and movement patterns to a greater extent to avoid jumping.

CHI-YUN KUO, GARY B. GILLIS and DUNCAN J. IRSCHICK (2012). Take this broken tail and learn to jump: the ability to recover from reduced in-air stability in tailless green anole lizards [Anolis carolinensis (Squamata: Dactyloidae)] Biological Journal of the Linnean Society DOI: 10.1111/j.1095-8312.2012.01958.x

Anolis Cuvieri Adventure

For many of us, the academic summer has finished or is ending imminently. In Boston, the temperature is falling, and most in the Boston area woke up to temperatures in the low 50s this morning. At this point, I thought the timing would be good to revisit (with some nostalgia) the manner in which I started the summer – with a three week field trip to Puerto Rico.

In June I was in Puerto Rico primarily to help my first Ph.D. student, Kristin, start her thesis project on urban ecology and adaptation in anoles. The focal species of Kristin’s research is the ubiquitous Anolis cristatellus, which, as anyone who has visited Puerto Rico will know, is equally common (if not more abundant) in heavily urbanized habitats as it is in natural forests. One species that is not found in urban areas, and, in fact, is fairly difficult to find in most habitats, is the Puerto Rican crown giant anole, Anolis cuvieri. We were lucky enough to see a few of these anyway, including one that I happen upon entirely by accident on the 60 acre finca where we stayed in a rental cottage for a little more than a week.

At night I was searching for invasive boa constrictors which are known from this part of the island, so as dusk approached I thought I’d try and take some photos of the sunset over the island’s western coast. Always on the lookout for A. cuvieri, I nonetheless somehow missed this individual in this pre-dusk shot (highlighted here by the red arrow). A perfect “find the anole” photo, but one in which I had initially “missed the anole” in spite of seeing it in person!

I initially missed this Puerto Rican crown giant, perched 20+ feet up a palm tree.

When I did spot him, he was far too high to capture with my meager 14 foot noose pole, so we just kept an eye on him. As the sun continued to set he did something interesting – he started to descend the trunk.

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