What’s In A Name?, Part II

Last week, I wrote a post on how the new classification for anoles proposed by Nicholson et al. 2012 might affect long term taxonomic stability for this group. That post generated some discussion, including, most recently, commentary by Kirsten Nicholson herself, explaining some of the reasons her group decided it was time to split up the anoles. Because that conversation is quickly becoming buried in the depths of Anole Annals, I wanted to continue it here on the main page. Also, I wanted to write some more to expand on some of the thoughts that prompted me to post last week, and I hope folks will continue to weigh in (note, the rest of this post will make a lot more sense if you first read my previous post, and the ensuing commentary). I’d like to make several general points:

1 – I think we can all agree that taxonomies should reflect accurate phylogenetic knowledge (I don’t think anyone here would disagree with this – let’s take it as a given that Linnean taxa should not be paraphyletic or polyphyletic).

2 – Beyond accuracy, I’m pretty sure there aren’t any rules governing the type or level of phylogenetic information that “should” be included in a binomial classification. Whether binomial names should reflect deep phylogenetic knowledge or shallow phylogenetic knowledge is a matter of opinion – I’d propose it’s completely subjective.

3 – The amount of phylogenetic information you can communicate in a binomial classification is trivial. It’s hard enough to represent phylogenetic structure across the depths of the entire Linnean hierarchy, and there’s almost no such information in Genus+species epithets. The goal of communicating finely resolved phylogenetic information probably should not be our main concern when dealing with genus names (so long as they are not phylogenetically inaccurate).

4 – Given that no one’s to say who’s right about what the appropriate phylogenetic scale of a genus is, and that Latin binomials are ineffective at communicating much of anything about phylogenetic information anyway, issues of stability are comparatively very important. It’s no small thing to propose a change for 88.6% (n = 343) of the scientific names of a group of species studied by thousands of people.

Since I think we can all agree that Linnean names should be applied solely to monophyletic groups, I’ll start with my second point, which is that there’s no “right amount” of phylogenetic knowledge that should be expressed in binomial names. Kirsten suggested we might all agree that “our classifications should mirror our phylogenetic knowledge.” I certainly agree with this statement in general, but I suspect I might disagree on some of the details. What sort of classification, exactly, would mirror our knowledge best? Should we assign genera to the smallest phylogenetic units about which we can be reasonably certain of monophyly, and continue to split them up into additional genera as we work towards resolving the entire bifurcating history of anoles? If so, we’ll probably eventually end up with a taxonomy that’s as finely parsed as that of the non-avian dinosaurs, where the genus:species ratio is 1:2 (I’m not even kidding – check it out…)! At this point, we’d have all sorts of cool binomials, like Nicholsonolis annectans and Mahlerolis gorgonae, but the genera would be functionally equivalent to species names (as they are in dinosaurs). This sort of reasoning (taxonomy should reflect ever-improving phylogenetic knowledge) is implied in the very title of the Nicholson paper, which seems to suggest that periodically, when the progress of systematics has advanced enough, “it is time” to reclassify everything (I think this contradicts the founding purpose of Linnean classification, but that’s another point). Anyhow, if this isn’t what it means for a taxonomy to mirror phylogenetic knowledge, then what exactly does that mean? Why 8 genera, and not 60, 16, or 4?

My main point here is that it’s a matter of opinion what kind of phylogenetic knowledge should be in a Latin name. One person might think that a genus should apply to the MRCA and all descendants of any two species similar enough to be confused by an experienced herpetologist (e.g., Anolis fraseri and A. biporcatus; see Williams 1966 for details). Another person might maintain that a genus should have 20 species max, no exceptions. Both are entirely matters of opinion, and such opinions abound when it comes to systematics. But since there are no official guidelines on the matter, I don’t think that such opinions can suffice to justify a disruptive taxonomic change.

I next wanted to criticize the logic of amending genus names to reflect systematic developments. The reasons are that (a) there’s very little phylogenetic information in Latin binomials, and (b) any change in the names of genera will result in a gain of some phylogenetic information (shallower information) at the expense of other phylogenetic information (the deeper stuff).

Linnean binomials contain next to no phylogenetic information. When we look at a list of scientific names, all we know is that congeners are more closely related to one another than they are to members of other genera, and other than that, they don’t tell us anything about phylogeny. To illustrate this, I created “binomial phylogenetic trees” for the Iguanidae (or Iguania, depending on who you follow..). I included all species in “Iguania” from the Reptile Database. Here’s what the traditional classification looks like, with Anolis highlighted in red:

Binomial “phylogenetic tree” of iguanian genera, following the traditional classification.

Continue reading

A History of Head Bobs

ResearchBlogging.orgIf you were to cast lizards as characters in a remake of The Great Gatsby, anoles, of course, would have to be cast as protagonist Jay Gatsby. “What Why Gatsby?” you might ask.  Well, like Gatsby, anoles have gaudy, aggressive displays that show fitness and define territory. Also like Gatsby, (SPOILER ALERT) anoles cannot seem to escape their past (for the headbob part of their display, at least).

In a recently published paper, Terry Ord investigated whether past ancestry explains variation in Anolis diplays better than evolution by natural selection. To begin to ask this question, Terry compiled data on the average time of headbob and dewlap bouts for species for a Western Caribbean Anolis lineage (Jamaica, Cayman Islands, Cuba) and for an Eastern Caribbean lineage (Puerto Rico and Hispaniola). The data and phylogenetic relationships are shown in Figure 1. Continue reading

Species Richness Patterns in Caribbean (and Mainland) Anolis V

In our comparison of mainland and island species turnover, we sampled mainland communities by randomly superimposing each of the Caribbean islands on the mainland five times. In each of these mainland island-shaped regions, we then tallied how many species were present and then measured turnover between regions.

ResearchBlogging.org

Oceanic islands are famous for their high endemism. We all love Darwin’s finches, Hawaiian honeycreepers, Komodo Dragons, and all those weird things on Australia. These endemic wonders suggest that islands may be home to unique processes of diversification.  However, there are many mainland regions that rival islands in their level of endemicity, especially in mountainous regions that have strong dispersal barriers and environmental gradients. Thus, you could ask, are islands truly special? Continue reading

High Flyin’ North Carolina Anoles

This NC green anole is on the ground, but mostly they apparently are up in the trees. Photo from http://www.wildherps.com/images/herps/standard/08041129PD_green_anole.jpg

We’ve had a lot of talk here on AA about how green anoles can be found low to the ground in places where A. sagrei has not invaded. But Manuel Leal says that’s not so in sagrei-less North Carolina, where all the ones he saw in the woods were high in the trees. What gives?

Anolis Electrum: The Amber Anole From Mexico

A recent photograph by Emma Sherratt. Amber tends to darken over time, which explains the deep orange color.

We’ve had previous posts on fossil anoles in amber. Emma Sherratt is currently studying them and has examined some three dozen specimens. All of these are from the Dominican Republic. Except the first one ever discovered, a Mexican piece described by Skip Lazell in 1965. Anolis electrum, as it was named, has had a pretty quiet scientific life. Now middle aged, the species has not been the subject of any subsequent research in the 47 years of its existence. But now it’s in the spotlight, as its phylogenetic position and dating may be pivotal for the recent calculation by Nicholson et al. that anoles originated more than 100 million years ago. In this post, I summarize what is known about A. electrum (examine the short original paper for yourself!). No doubt, we’ll be hearing more soon about the relevance of this species–specifically its phylogenetic placement and age–for dating anole diversification.

As you can see for yourself in the photo above, there are actually two pieces, a front half of a lizard and a back half a lizard. Since they were found together (or at least made it to the Paleontology Museum at UC-Berkeley together) and are matching in size, it seems like too much of a coincidence for them not to come from the same animal. Various aspects of the animal’s scalation are discernible, including some nicely visible toepads. Lazell stated that all that was left was skin, or the impression of skin, the bones having been eaten away, but Emma’s cat scanning has shown that this is not quite correct (see below).

Based on the specimen, what can be said about its phylogenetic placement? All anoles in Mexico today are from the Norops clade. Unfortunately, the primary character for identifying Norops is the shape of the caudal vertebrae, which cannot be discerned in this tailless specimen. Lazell compared the scalation of this specimen (a 26 mm juvenile) to various species, and found that the scalation was unlike most species. He concluded that electrum was most similar in scalation to A. fuscoauratus, A. maculiventris, and A. chloris, and among species found in Mexico, to A. limifrons (full quotations at the bottom of this post).

What should we make of all of this? It’s important to remember that this paper was published in 1965, prior to the description of many extant anole species and a year before Willi Hennig’s classic introduction to cladistic analysis was translated into English. This is a purely phenetic comparison of the amber baby lizard to known species, clearly non-phylogenetic and utilizing characters that now are recognized to generally have little higher level systematic utility in anoles. And the conclusion is that it is either a Norops clade anole (fuscoauratus, limifrons or maculiventris) or a Dactyloa species (chloris).

The other question one might have is: how old is this fossil? Dating amber is notoriously difficult. Solórzano Kraemer reviewed all of the data on Mexican amber bearing deposits in the 2010 volume Biodiversity of Fossils in Amber from the Major World Deposits and concluded: “In summary, it can be said that Mexican amber can be correlated with Dominican amber, with an age of approximately 15-20 million-years-old.” In other words, Mexican and Dominican amber anoles were contemporaneous.

Did anyone notice anything odd on the fossil of the lizard posterior (B, above)? Continue reading

2012 Anole Photo Contest: Last Chance To Enter

We’ve already received a lot of great entries, but there’s room for more! We’re reprinting the announcement below. Deadline is September 30, so submit today!

Last year we had an Anolis photo contest and produced a 2012 anole calendar. Both were wildly successful. Today, Anole Annals is pleased to announce it is combining both. We herewith announce the 2012 Anole Photo Contest. The goal of the contest is to identify 12 winning photos.  The grand prize winner will have her/his photo featured on the front cover of the 2013 Anole Annals calendar and will receive an autographed copy of Karen Cusick’s lovely book, Lizards on the Fence. The second place winner will receive a copy of the calendar and have her/his photo featured on the backcover of the calendar.

The rules: please submit photos as attachments to anoleannals@gmail.com. To ensure that submissions with large attachments arrive, it’s a good idea to send an accompanying e-mail without any attachments that seeks confirmation of the photos receipt.  Photos must be at least 150 dpi and print to a size of 11 x 17 inches. If you do not have experience resizing and color-correcting your images, the simplest thing to do is to submit the raw image files produced by your digital camera (or, for the luddites, a high quality digital scan of a printed image). If you elect to alter your own images, don’t forget that its always better to resize than to resample. Images with watermarks or other digital alterations that extend beyond color correction, sharpening and other basic editing will not be accepted. We are not going to deal with formal copyright law and ask only your permission to use your image for the calendar and related content on Anole Annals. We, in turn, agree that your images will never be used without attribution and that we will not profit financially from their use (nobody is going to make any money from the sale of these calendars because they’ll be available directly from the vendor).

Please provide a short description of the photo that includes: (1) the species name, (2) the location where the photo was taken, and (3) any other relevant information. Twelve winning photos will be selected by readers of Anole Annals from a set of 28 finalists chosen by the editors of Anole Annals.  The grand prize winning and runner-up photos will be chosen by a panel of anole photography experts. Deadline for submission is September 30, 2012.

Lacertid Pays The Price For Being Mistaken For A Gecko; Thanks Geico

We’ve previously discussed cases of anoles being mistaken for geckos, as well as the very  negative effects that cats can have on green anoles. Turns out that anoles aren’t the only lizards that serve as gecko doppelgangers. And you might think that people wouldn’t mistake lacertids for anoles, but apparently that happens, too.

More On Nicholson et al. 2012: Let’s Look At Their Methodology

ResearchBlogging.orgMost people who have commented on the blog about Nicholson et al. 2012 have focused on whether is it really necessary to name all these inferred clades as genera. I agree with those who state it is completely unnecessary and disruptive, and that there are alternative ways (e.g., assigning names to relevant clades independent of the genus rank) to describe the diversity of Anolis. That said, I would like to direct the discussion towards the methodology used. Yes, there are a lot of missing ND2 data in their dataset (e.g., all of the new data presented in Castañeda and de Queiroz 2011 is missing), but I think it is more relevant to consider how they treated the data they did include. First, the molecular partition of their DNA: the protein coding gene ND2 was not partitioned into codon positions, which has been shown to be the best strategy (e.g., Schulte and de Queiroz, 2008; Torres-Carvajal and de Queiroz, 2009; Castañeda and de Queiroz, 2011), and instead, they chose to set a different partition for each of the tRNAs included (five) and one more for the origin for the light strand replication piece (which is ~30 bases long). As the Bayesian analysis requires a large-enough number of characters to estimate the parameter values for the model selected, I thought it was recommended to have partitions of more than ~300 bases (and I can’t think from the top of my head for a specific citation here). Neither the OL nor any of the tRNAs is close to this size (and the AICc, the corrected Akaike Information Criterion, intended for small sample sizes should have been used to select the best fitting model here instead of the regular AIC).(For more on partition selection and consequences of under– or overparameterization, check Brown and Lemmon, 2007 and Li et al. 2007). This should raise an eyebrow about the thoroughness of the analyses. However, in reality, I think this would have little effect on the actual phylogeny. Those clades that are strongly supported would be robust enough to withstand model and partition misspecifications.

On the other hand, the treatment of the morphological characters might have more serious effects on the resulting topology. Nicholson et al. explain that they used Poe’s 2004 morphological data as is, but without the complex coding system he used for continuous and polymorphic characters, and instead considering all possible characters to be equally weighted. (To be fair, Poe did use equal weighting for characters in his analyses; the cost of changes between states within a single character is what is different). Poe coded continuous characters using a gap-weighting method, which divides the range of a continuous character into discrete segments, maintaining information on the order of the character states and the magnitude of the difference between them, and he coded polymorphic characters using a frequency method, which keeps track of the fraction of individuals within the sample that shows a given state. From what I understood, Nicholson et al. considered all changes to be of equal cost, so transitioning from the smallest head to the largest head, or from having all individuals showing condition x to all individuals showing condition y (where some taxa exhibit both conditions), will cost 26 steps, which is the cost of changing from state a to state z (as recognized by Poe). This means, in the combined parsimony analysis, a transition between the two extreme states in a continuous or polymorphic morphological character is equivalent to [single] DNA substitutions at 26 different positions [characters]. Moreover, changes in those morphological characters that were not continuous or polymorphic would cost only a few steps. This weighting scheme (in the parsimony context) will actually give a higher weight to some morphological characters, which is exactly the opposite of what the authors were aiming for (i.e., equal weights). The effects of this unbalanced weighting on the resulting topology? Not sure, but I’m going to guess not insignificant!

One last thing. Several of their proposed genera (Dactyloa, Deiroptyx, Chamaelinorops and Xiphosurus) are not monophyletic on their combined data tree, the one that supposedly serves as the basis for their taxonomy…

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

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. Continue reading

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

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.

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,

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

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. Continue reading

Cuban Owls Eat Big Anoles – New Research by Yudisleidy López Ricardo

Here on the Anole Annals we like to talk food. Although anoles are predominantly insectivorous creatures, we have documented some of their stranger eating habits on this blog. For example, through recent research we have learned that they are more frugivorous than previously thought. They also include other vertebrates into their diets, such as frogs. Chamaeleolis anoles, we have learned, have specialized molars to aid in crunching mollusks.

Sadly, however, anoles are often also on the receiving side of predation. Anoles are important prey items for many different animals. Sometimes, even plants get their fill on anoles.

In her recently published undergraduate thesis, Dr. Yudisleidy López Ricardo from the University of La Habana, Cuba discusses the diet of the barn owl (Tyto alba furcata) in several localities in Villa Clara and Ciego de Ávila. Dr. López Ricardo examined nearly 300 owl pellets (regurgitated bits that contain food remains) and found 69 different prey types. As expected, small mammals such as the house mouse and black rat were common prey items. A novel finding of this study, however, is that large species of anoles, namely A. equestris, A. porcatus, and even Chamaeleolis sp. lizards were found in the owl pellets. Smaller anoles, including A. jubar, A. sagrei, and A. lucius were also found in the diets of the barn owl. The authors also found that a different herp, the Cuban tree frog, Osteopilus septentrionalis, was not uncommonly found in owl pellets, but this species is nocturnal.

The finding that anoles are a small, but important, component of this species’ diet is quite interesting in light of the fact that Tyto alba, like most owls, is nocturnal. The main question for me is how they are finding and catching anoles. Owls rely heavily on sensitive hearing to locate moving prey at even great distances. But anoles are predominantly diurnal creatures, and are typically asleep and quite still by nightfall. Owls also have great vision and may be spotting anoles during crepuscular hours. Or are they opportunistically feeding on anoles? Perhaps a different predator scares an anole out of its sleeping site and owls are snatching up fleeing anoles.

Any thoughts from the Anolis community on this interesting finding?