The Origins of Anolis carolinensis

Fig. 1: Figures illustrating Cuban origins of A. carolinensis from our 2005 paper in Molecular Ecology. Green shading indicates the range and phylogenetic position of A. carolinensis, blue shading indicates Cuban populations related to A. carolinensis. The arrows indicate possible dispersals from Cuba, some of which are supported by phylogenies (including the dispersal from Cuba to the continental United States indicated by the bold arrow).

With all this discussion of the green anole’s genome, it seems like a good time to remind everyone of how Anolis carolinesis came to be the model organism that it is today.  The simple answer, of course, is that A. carolinensis is the only species of anole endemic to the continental United States.  As such, its always been the anole species most accessible to the broadest range of researchers.  The deeper answer – and the focus of this post – concerns how A. carolinesis happened to become the continental United States’s only native anole in the first place.

Anole Annals: Your One Stop Anole Genome Information Source

For information on why the anole genome is useful for evolutionary studies, go here.

For information on how the genome is already being used in research, try here, here, here, here and here.

For the history of discovery and study of anoles, go here.

For the evolutionary history of the green anole, check this one out.

For a great story, don’t miss this one.

For great pictures of anoles and their dewlaps, try here, here, and here  (among others).

For many other topics in anole ecology, behavior, and diversity, try looking up terms in the blog’s search window.

What’s The Anole Genome Good For?

One of these species has had its genome sequenced, and the other has independently evolved to look very similar and live in the same environment. The anole genome will make anoles an even more powerful group in which to study evolutionary convergence. Photos by Melissa Losos (left) and Pete Humphrey (right).

When the genome of Anolis carolinensis is finally published, most attention will focus on how this genome, the first reptile to be sequenced (not including birds), differs from other vertebrate genomes, and what these differences may tell us about genome evolution. No doubt this will be interesting, but the real value of this genome–in my unbiased opinion–resides in the questions we finally will be able to address about the evolutionary process, particularly in one model system of evolutionary study, Anolis lizards. Chris Schneider published a perceptive article, “Exploiting genomic resources in studies of speciation and adaptive radiation of lizards in the genus Anolis,” on this topic three years ago, and I will briefly expand on his points here.

An anole genome will be useful for evolutionary studies in two ways. First, a long-standing question in evolutionary biology concerns the genetic basis of convergent evolution (i.e., when two or more evolutionary lineages independently evolve similar features). Do convergent phenotypes arise by convergent evolution of the same genetic changes, or do different lineages utilize different mutations to produce the same phenotype? In other words, does convergence at the phenotypic level result from convergent change at the genetic level, or can different genetic changes produce the same phenotypic response? In the last few years, molecular evolutionary biologists have produced a wealth of studies investigating whether convergent changes in coat color in rodents, eye and spine loss in fish, bristle loss in fruit flies and many other changes are the result of changes in the same gene, even some times by the very same genetic mutation. Underlying these questions are more fundamental questions about constraints and the predictability of evolution (these topics have been reviewed a number of times in the last couple of years, most recently in a paper by me, in a paper which refers to other recent reviews).

The anole ecomorphs, habitat specialists behaviorally and morphologically adapted to use different parts of the environment. The same set of ecomorphs (with several exceptions) have evolved independently on each island in the Greater Antilles. Figure from "Lizards in an Evolutionary Tree," based on earlier figures in Ernest Williams' work.

Anolis lizards are, of course, the poster child for evolutionary studies of convergent evolution. Indeed, convergence has run rampant in this clade. AA has prattled on endlessly about the famous anole ecomorphs, a set of habitat specialist types that have evolved repeatedly on each island in the Greater Antilles to occupy different habitat niches. This convergence is usually studied in terms of limb length, tail length, and toepad dimensions: arboreal species have big toepads, twig species short legs, grass species long tails, and so on, with these traits independently evolving many times. But the ecomorphs are convergent in many other traits that have received less attention: head and pelvis dimensions, sexual dimorphism in both size and shape, territorial and foraging behavior, to name a few, and the more closely we look, the more convergent traits we find. And, further, anole convergence is not entirely an ecomorph phenomenon; some traits vary within an ecomorph class, but are convergent among species in different ecomorph classes, for example, thermal physiology and dewlap color.

In other words, there’s more convergence in Anolis than you can shake a stick at, and the availability of the anole genome sequence will provide the tools to investigate its underlying genetic basis.

Anolis Equestris On The Big Screen

ok, the little screen.

 

Anolis Roquet Males Displaying On Exhibit

Acrobatic bushmaster says hello!

At the Nashville Zoo we have a large mixed species exhibit that contains two species of dart frog, a bushmaster, Gonatodes, and 2.3 Roquet’s anoles (Savannah anoles).  We are working on some new graphics where we hope to incorporate videos, and in that attempt we captured some great footage of our two male A. roquet displaying towards one another.  The two males are approximately a foot apart and right in front of the glass of the exhibit.  The females were watching anxiously as shortly after the video stops one male chases the other up a tree.  I hope you enjoy the clips.

http://www.flickr.com/photos/66876684@N04/6086156957/in/photostream

http://www.flickr.com/photos/66876684@N04/6086649388/in/photostream/

A Brief History of Anoles in Research

The sequencing of the genome of the  green anole (Anolis carolinensis) is a landmark in the age of genomics, and a highpoint in the annals of anole studies.  It is the first complete genome sequence of a reptile, and a great step forward in the development of comparative genomics. Results are already coming in: Matthew Fujita, Scott Edwards and Chris Ponting have a paper in press in Genome Biology and Evolution, using an earlier release of the anole genome, showing that the green anole genome lacks the large guanine-cytosine rich regions (called isochores) that are characteristic of birds and mammals. Is this lack unique to the green anole, or a feature of some larger group? We don’t know, of course, because the green anole is the first non-avian/non-mammalian amniote to be sequenced, but it is a hint of new things to come. As Rich Glor put it in his commentary here at Anole Annals,

Fujita et al.’s work  is a good example of the insight offered by comparative genome sequencing; as the number of available genomes expands, this work is sure to continue to challenge overly simplistic assumptions about genome architecture and evolution derived from biased sampling of the tree of life.

Green Anole, Anolis carolinensis, New Orleans, LA. Note the expanded subdigital toepads.

So why was an anole chosen to be the first reptile sequenced? The short answer is that, among a small group of candidate species of reptiles announced by the NIH in May of 2005, the response of the biological community favored the green anole over the garter snake. But the longer answer includes why the community preferred the green anole: it has long been the subject of diverse physiological, behavioral, ecological, and other  studies (a “model” organism in the functional biology sense), and, more importantly, anoles in general have been favored and favorable materials for a diverse array of physiological, behavioral, ecological, evolutionary, and zoogeographic studies over many decades. The goal of this post is to review a bit of the rich history of anole studies to provide some background on why so many researchers have found anoles to be vastly interesting animals.

Does Global Climate Change Threaten Tropical Lizards?

Anolis allisoni (photo from http://www.kingsnake.com/westindian/anolisallisoni2.JPG)

Everyone’s worried about global warming. For a long time, frogs hogged the herpetological spotlight, with concern that the global amphibian crisis might be driven by climate change. However, in recent years, there has been a growing realization that lizards may be in trouble, too, and again the finger has been pointed at climate change.

One hypothesis put forward by Ray Huey and colleagues is that as temperatures warm, open-adapted species will be able to invade forests, which previously had been too cool for them, and the cool-adapted forest lizards, living in a now warmer home and faced with competition from the invaders, would have nowhere to go and would be in big trouble.  Preliminary data from Puerto Rico support this model, and Huey and colleagues have returned to the enchanted island to further test the hypothesis.

Michael Logan, a graduate student at Dartmouth, has set out to test this idea elsewhere, working in the Bay Islands off the coast of Honduras. These islands are particularly interesting because they are one of the few places where Caribbean and mainland anole faunas meet, with members of the sagrei and carolinensis species groups of Cuba coexisting with several mainland species. This juxtaposition is interesting in its own right, but it turns out that the Caribbean species are warm-adapted, open-living species, whereas the mainland species are cool-adapted, forest types. Logan’s goal is to test the hypothesis that as warming occurs, the warm-adapted species will be able to enter the forest, with potentially adverse effects on the species therein. In a recent issue of Biodiversity Science, a newsletter put out by Operation Wallacea, Logan reports preliminary results from last year’s field season, and they’re not what you might expect.

Irene Passing Over Abaco

According to NOAA, the eye is over Abaco right now, with sustained wind speeds of 115 mph

The Impending Armageddon II

Doesn’t look good. Staniel and particularly Abaco are right in the path of Irene–Category III for Abaco. Hang in there, lizards and people!

Anoles: They’re Just Like Us!

a juvenile Anolis orcesi, in the afternoon of an all-day recording

They wake up, but hit the snooze button before they really get moving.  They poop, grab something to eat, and then check out the neighborhood.  They take wrong turns, and have to turn around when they reach a dead end.  Young boys try to impress each other with their dewlaps.  They take naps in the afternoon, and yawn throughout the day. They even sleep in the same bed, most nights.

As Jonathan Losos hinted at in an earlier post, we observed 4 Anolis orcesi individuals from dawn to dusk (12 hours!), and several more individuals for 1 to 6 hours, in the vicinity of Baeza, Ecuador.  More to come after several months of video analysis!

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