All posts by Jonathan Losos

About Jonathan Losos

Professor and Curator of Herpetology at the Museum of Comparative Zoology at Harvard University. I've spent my entire professional career studying anoles and have discovered that the more I learn about anoles, the more I realize I don't know.

What You Can Learn from Watching a Lizard for a Long Time

Photo by Ambika Kamath

Behavioral research is often reduced to a large set of data points, necessary of course for statistical analysis. But sometimes what gets lost is actual knowledge of what animals do in their natural habitats. There’s no substitute for just watching an animal over the course of a day or  a week. Often what you’ll see is that animals are not little automatons, repetitively undertaking particular actions in accord (or not) with our theories. Rather, they have lives where they do all kinds of idiosyncratic behaviors, the sort of quirky detail that often get lost in high-falutin’ analyses of behavior. Ambika Kamath demonstrated just this in her recent post, “A Week in the Life of U131.” Here’s the first paragraph. You’ll have to go to her website to read the rest:

When you’re collecting data on the behaviour of individual animals over time, as I am this summer, your observations sometimes feel less like a collection of numbers and more like a collection of personal narratives. Of course, the data are both numbers and narratives, and when it comes time to analyze this collection of datapoints and understand the patterns that emerge from it, the numbers will be all that matter. But in the meanwhile, before I can look the bigger picture, I enjoy considering the individual narratives. And this week, I encountered a lizard whose story illustrates why it’s worth considering these narratives at all.

Anole Annals Wants You…to Write a Post!

The goal of Anole Annals is to be the clearinghouse for all things Anolis, the place that the anole community turns to for the exchange of information or ideas. To do so, we welcome–no, heartily encourage–contributions from anyone and everyone.

Who can post? Anyone who has something to say about the biology, natural history, or amazing-ness of anoles (well, within reason–we leave anole husbandry and sales issues to other websites). And fear not–you’ll have an audience. Anole Annals is now routinely visited by 600-1000 readers a day. That sounds like a broader impact to me!

Anole Annals is a good place to let the anole community know what you’re working on, like Ambika Kamath’s recent overview of her fascinating work on anole social behavior. And, it’s a great way to spread word of your recently published work–why not provide a short precis or tell the backstory of how the paper came to be, like Liam Revell recently did? It’s a great way of giving people the short story of what you’ve done and get them interested in reading the whole paper.

If you are fortunate to live in an anole-inhabited region, tell us about your local species, like David Alfonso’s recent post on the anoles of Colombia. And if you’ve observed something unusual, here’s a good place to report it, like Graham Reynolds note on twig anoles using mangroves.

And it’s just a great place to ask a question, post a photo, or report an observation. Plus, announcements of relevant conferences or personal milestones, such as newly-minted Ph.D.s, are always appropriate.

Posting is easy, and really doesn’t take much time. More than 100 scientists and anole enthusiasts have written posts–you should too! And if you’ve done so before, you’re overdue for another one. Don’t overthink it–just post today!

Clay Models with Dewlaps Are Attacked More than Clay Models Without Them

modelIn recent years, biologists have put out clay model lizards to measure attack rates by birds and other predators. In a recent study published in Herpetology Notes, Vazquez and Hilje put a twist into this approach, by ornamenting some faux lizards with a colorful dewlap-like structure. Sure enough, those models were attacked more. Here’s their abstract:

We investigated how predatory attacks on Norops lizards occur in old-growth and secondary pre-montane wet forests in Costa Rica using clay models. Using models with secondary sexual characteristics also permitted comparison of attack rates between males and females, as well as the specific site of the attack (head, body, or tail). Birds were found to account for the bulk of predation attempts on the lizard models in both forest types, and there was no significant difference in attack frequency between forests. Attempts were more frequently made on males, indicating that colorful, bright dewlaps used to attract females and for territorial displays might also function in attracting predators. Male models received more attacks on the head, suggesting that birds may preferentially target areas of brighter coloration, whereas female models received more attacks on their tail, indicating that birds may not have adopted any avoidance of a potentially less profitable region from which tails can be autotomized.

Lizard Systematics: Morphology and Molecules Redux

Closely related to snakes after all?

Two years ago, I wrote an AA post on lizard phylogenetics, summarizing the results of a paper by Gauthier et al. that claimed that analyses based on morphological and molecular data produced very different phylogenies. Moreover, Gauthier et al. argued that the morphological data provided absolutely no support for the phylogeny suggested by the molecular data.

The issue has now been further considered in a recent paper by Reeder et al. in PLoS One. These authors provide some new morphological data and add a tad of previously unutilized molecular data; with these data sets, they recover essentially the same disagreement in phylogenies. However, what is new in this paper is that they perform a combined analysis that analyzes morphological and molecular data together. The results of this analysis are firmly in agreement with the molecular data. To address the possibility that this is simply a result of the much greater quantity of molecular characters, the authors also conducted analyses that essentially weighted the molecular and morphological data equally. Still, the result was the tree produced by the molecular data alone.

Perhaps the most striking point in the Gauthier et al. paper was the claim that the morphological data gave absolutely no support for the molecular tree. This suggested, in turn, that if the molecular tree is correct, then morphological evolution has been extraordinarily homoplasious. However, Reeder et al. dispute this claim, finding that a number of morphological characters support the molecular phylogeny.

Reeder et al. also broke the morphological characters into six subsets: cranial characters; characters related to the jaws, teeth, and hyobranchial apparatus; characters related to the vertebral column; other postcranial osteological characters, mostly related to the limbs and limb girdles; miscellaneous morphological characters, including morphology of the osteoderms, scleral ossicles and tongue; and characters of squamation and external morphology. Their analysis found that only the cranial characters were incongruent with the molecular phylogeny, and they suggested that these were the characters in which homoplasy was rampant, leading to false signal in the morphological analyses.

Overall, the authors make a strong case that the molecular phylogeny is likely to be the correct one and that morphological data, particularly cranial characters, are misleading due to homoplasy. It will be interesting to see whether and Gauthier et al. respond to these analyses.

Here is the take home message from the discussion of Reeder et al.’s paper:

Our combined analyses strongly suggest that the phylogenetic hypothesis for living squamates based on the molecular data is correct. Specifically, our results support the hypothesis that Iguania is placed with snakes and anguimorphs, and not at the squamate root (as suggested by morphological data alone). Our conclusions are based on several lines of evidence, including:

(a) combined analyses of the relevant molecular and morphological data supports the molecular placement of Iguania, even when the molecular dataset is reduced to only 63 characters, less than one tenth the size of the morphological dataset,

(b) mapping morphological characters on the combined-data tree shows that there is actually hidden support for the molecular hypothesis in the morphological data (similar to the number of characters supporting the morphological


(c) the morphological dataset is dominated by misleading phylogenetic signal associated with convergent evolution of a burrowing lifestyle and associated traits, and a similar problem associated with feeding modes may explain the morphological placement of Iguania, and

(d) the morphological hypothesis is unambiguously supported by only one of six subsets of the morphological data. Conversely, we find no evidence for hidden signal supporting the morphological hypothesis among the 46 genes in the molecular dataset; no genes support this hypothesis. Further, the failure of some genes to fully support the molecular placement of iguanians in Toxicofera seems to be associated with sampling error (i.e. shorter genes).

Update on the Anoles of Singapore

Anolis sagrei displaying in front of a supertree at the Gardens by the Bay

Anolis sagrei displaying in front of a supertree at the Gardens by the Bay

Two years ago, we posted on a paper in Nature in Singapore documenting the occurrence of the festive anole, Anolis sagrei, in Singapore. The ubiquitous colonizers had turned up in the newly created Gardens by the Bay, an enormous new botanical garden built on reclaimed land at the southern end of the island. AA decided to look into the situation further and sent this correspondent to the “Lion City” to report back on the situation.

Reporting for duty at the Gardens at approximately 930 am on the morning of 16 April, we quickly determined that the lizards are common in the lushly planted gardens pretty much wherever we went. The only exception was an open meadow housing a statue, where we did see an introduced agamid lizard (below). This area was only searched for three minutes, however, and it would not be surprising to find the anoles in the shrubbery surrounding the statue (below). The other place the lizards were not seen were in the two spectacular indoor cool houses, the Flower Dome and the Cloud Forest. Both are kept at temperatures possibly too low for the lizards, and also likely are fumigated.


Introduced Calotes versicolor at Gardens by the Bay

IMG_7370xOtherwise, however, the anoles seem to be everywhere and it seems unlikely, given the luxuriance of the vegetation, that they can be eradicated (and we know how well such elimination efforts have fared in Taiwan—not).

The question is whether the anole will spread to the rest of Singapore, and from thence to Malaysia. Given the heavy motor traffic bringing visitors to the gardens, it seems inevitable that the anoles will hitch-hike their way across the bridge and colonize the main island of Singapore, which is for the most part one big Anolis sagrei habitat, with plenty of tropical vegetation everywhere. Moreover, the gift shop at the Gardens was selling orchids. I don’t know where they came from, but if on site, that is a great way to distribute brown anoles far and wide.

IMG_2609xIt’s not clear whether the anole is already present on the Singaporean main island. One commenter on our previous post said that it had been seen elsewhere, and I was told that there were unsubstantiated reports that it had been observed in the Singapore Botanical Garden. I spent several hours there myself and saw the anole mimic pictured on the right, but no anoles.

My prediction is that in ten years, Anolis sagrei will be very common in Singapore. But let’s see what the varanids, the Calotes and the birds have to say about that.

New Monograph on the Reptiles and Amphibians of Southern Florida


Meshaka and Layne have just published a masterful review of the native herps of southern Florida in Herpetological Conservation and Biology (freely downloadable). Of most interest to readers of these pages is the treatment of Anolis carolinensis, and it is indicative of the quality of this work: the six pages devoted to the natural history of the green anole is the most authoritative and comprehensive of which I’m aware, covering the literature for this species not only in southern Florida, but throughout its range. This monograph is the starting point for anyone interested in green anole biology. In addition, this section shows how surprisingly little we know about the biology of this species. For example, most of the information on green anole diet comes from Wayne King’s work from 1966.

This volume will be useful to anyone interested in the herpetology of southern Florida.

The Mystery of the Beat Up, Passive Crested Anole

Photo by Janson Jones

Look at this poor fellow. Over on Phostracks, Janson Jones describes him and his demeanor thusly:

“Still, the most compelling characteristic of this anole was its passiveness. It wasn’t dead. Far from it, actually. The anole actively watched me, tracking me with its eyes. Still, when I rolled the lizard on its back, it just laid there like a puppy wanting a belly rub. Except for two or three miserable feeble attempts at escape, this anole was entirely passive (and clearly in need of a good dinner). It was just worn the hell out and in dire need of a vacation.

I placed the anole on the side of an unpopulated tree trunk to see if it could or would hang on (see image bottom left; image is rotated 90 degrees). No problem. It did just fine. It even moved around a bit. Then, when I started creeping back close again, nothing. It just sat there. Passive.

So, to quote Diondre Cole, “What’s up with that?” Any ideas? Any opinions? Any speculations?”

For more photos and entertaining description, check out the full post.

The World’s Most Beautiful Anole? Anolis equestris potior

equestris potior

We’ve had posts on this spectacular anole before. Jesus Reina Carvajal reports: “During the last three years, I have been lucky to see them every single time I have visited the area but they are really hard to find. I find them in the wild on Cayo Santa María. They live exclusively on that island. Nowhere else in the world.

They eat insects, other lizards and small birds. They have a powerful bite. This last individual I could follow during hours since I saw it early in the morning and I could make many pictures and films until it left the place. That day I felt very happy!”

More of Jesus’s photos can be seen in the Flickr album he created.

Photo by Jesus Reina Carvajal

Photo by Jesus Reina Carvajal

Photo by Jesus Reina Carvajal

Photo by Jesus Reina Carvajal

Grey-Dewlapped Crested Anole

grey crested

Stroud and Beckles published this photo of a crested anole with a half-grey dewlap in the December 2014 issue of Herpetological Review. Cresteds dewlaps are usually orange or yellow–this is very unusual.

This is reminiscent of the famous gray-dewlapped A. carolinensis as well as this odd crested anole found by Neil Losin.

What’s up with these wacky anoles?

Placing Extinct Species in a Molecular Phylogeny Using Quantitative Characters: A Case Study Using Anolis roosevelti

Liam Revell writes:

My co-authors (Luke Mahler, Graham Reynolds, & Graham Slater) and I recently presented a ‘new’ method for placing recently extinct taxa into a backbone molecular phylogeny on the basis of quantitative trait data. I say ‘new’ with quotes, because our methods derives closely, with full credit given where due, from a Maximum Likelihood phylogeny inference approach presented originally by Felsenstein (1981, 2002).

The idea is basically as follows. We start with a time-calibrated molecular phylogeny containing N – 1 species, and a single taxon of interest (the Nth taxon) whose placement in the tree is of interest, but for which molecular characters are missing. If we have quantitative trait data from one or more characters for all N species in the tree, we can use an approach based on Felsenstein (1981) to add this taxon to our base phylogeny using the statistical criterion of Maximum Likelihood.Revell_etal.Figure3_1In our article (Revell et al. 2015), we demonstrate that the method works pretty well in theory. In fact, for more than a few quantitative characters & particularly for trees of large size, the method often places the missing taxon in our dataset in a topological position that is identical to its true position. (See figure below, reproduced from our article.) In the figure, white bars show the performance of our method (compared to grey bars which represent placement at random). In all cases, lower values indicate that the estimated tree is closer to the generating tree.

The question you’re probably asking yourself (and quite rightly so) is: what could this possibly have to do with anoles? The answer is that we applied the method to the unusual case of Anolis roosevelti. Anolis roosevelti, as many readers of this blog likely already know, is a mysterious crown-giant anole from Culebra and (probably) the Spanish, U.S., and British Virgin Islands, excluding St. Croix. It is only known from a few specimens and was last collected in 1932. Aside from some unconfirmed reports, it has neither been seen nor heard from since. Unfortunately – and tragically given the impressive nature of this creature – all but the most optimistic anole biologists agree that this species is most likely extinct. (Many of us, the author included, still holds out hope, of course.) The figure below shows the type specimen of this impressive creature. (Figure from our article and image courtesy of the Museum of Comparative Zoology at Harvard.)

figure2Since no prior investigator has collected molecular characters from this species, and the prospects for so doing in the future are somewhat mixed (for reasons that we explain in the article), we thought Anolis roosevelti would represent an interesting test case for our method. Would A. roosevelti, we asked ourselves, fall out as sister to the Puerto Rican crown-giant, Anolis cuvieri, as sister-to or nested-within the rest of Puerto Rican anoles, or in another part of the tree entirely? Continue reading

Identify These Puerto Rican Anoles

Bill Schlesinger, one of the world’s most eminent biogeochemists and President Emeritus of the Cary Institute of Ecosystems Studies, turns out to have an eagle eye for anoles. While on a birding trip to Puerto Rico, his wife, writer Lisa Dellwo, snapped the photos below in the rainforest in the west central part of the island. Which species are they?

Lizards Falling out of the Trees in the Rainforest


Piero Angeli Ruschi from the Ornithology Department in the National Museum in Rio de Janeiro sent the pictures of Anolis punctatus shown here. His story:


It’s raining lizards!

“These are the pictures that I took of one of the Anolis specimens that had fallen that day. They fell from the canopy straight to open ground over an area of ~70 squared meters. About 10 specimens fell within ~3 min. It happened in Santa Lucia Ecological Station, at Santa Teresa, ES, Brazil in late september 2007 during the afternoom while I monitored a woodpecker nest.

The individuals were all the same as the one in that picture…green with a yellow circle around the eye…Those pictures might even include more than one individual—I am not sure if the one photographed on the ground was the same I captured (they stayed knocked out for a minute or so before running from where they landed).


I wish I had more data to publish a communication about it, as well as more evidence to explain the cause of it. My ornithological background leads me to the alternative hypothesis of some sort of “arena,” but I have no idea if such thing can be expected for these lizards.”

Piero Angeli Ruschi in his day jobobserving birds

Piero Angeli Ruschi in his day job observing birds


The Social Life of Lizards Revealed: Lizard Social Behavior Research in Gainesville

If you’re in Gainesville and come across a site like this, odds are you’re not at a crime scene, but rather Ambika Kamath’s study site, where she’s investigating the social organization of brown anoles. The standard view of anole social structure is that a male brown anole defends a large territory, excluding all other males and courting the several females that reside therein. Ambika’s work to date suggests that reality is a lot more complicated. Ambika provides an in-depth discussion of what she’s doing and why on her blog .

Cellular Mechanism of Tail Regeneration in Anolis carolinensis

Green anole with a regrowing tail. Photo from Daffodil’s Photo Blog.

Lorenzo Alibardi is conducting detailed studies of what the cells are doing as the tail regenerates. His latest work is now available online in the Journal of Morphology. Here’s the abstract:

Using an antibody against a lizard telomerase-1 component the presence of telomerase has been detected in regenerating lizard tails where numerous cells are proliferating. Immunoblots showed telomerase positive bands at 75–80 kDa in normal tissues and at 50, 75, and 90 kDa in those regenerating. Immunofluorescence and ultrastructural immunolocalization showed telomerase-immunoreactivity in sparCe (few/diluted) mesenchymal cells of the blastema, early regenerating muscles, perichondrium of the cartilaginous tube, ependyma of the spinal cord, and in the regenerating epidermis. Clusters of gold particles were detected in condensing chromosomes of few mesenchymal and epithelial cells in the regenerating tail, but a low to undetectable labeling in interphase cells. Telomeraseimmunoreactivity was intense in the nucleus and sparCe (few/diluted) in the cytoplasm of spermatogonia and spermatocytes and drastically decreased in early spermatids where some nuclear labeling remains. Some intense immunoreactivity was seen in few cells near the basal membrane of intestinal enterocytes or in leukocytes (likely lymphocytes) of the intestine mucosa. In spermatogonia, spermatids and in enterocytes part of the nuclear labeling formed cluster of gold particles in dense areas identified as Cajal Bodies, suggesting that telomerase is a marker for these stem cells. This therefore suggests that all of the sparCe (few/diluted) telomerase positive cells detected in the regenerating tail may represent sparCe (few/diluted) stem cells localized in regenerating tissues where transit amplifying cells are instead preponderant to allow for tail growth. This observation supports previous studies indicating that few stem cells are present in the stump after tail amputation and give rise to transit amplifying cells for tail regeneration.

A Four-Tailed Brown Anole

Four-tailed Anolis sagrei. Photo by Lynne Carpenter Ingram.

Four-tailed Anolis sagrei. Photo by Lynne Carpenter Ingram.

Photo by Lynne Carpenter Ingram

Photo by Lynne Carpenter Ingram

Lynne Carpenter Ingram took this photo of a quadricolous (I made that word up) brown anole. Here’s what she had to say on her Facebook page: “Last Sept I posted some pictures of a lizard I have living in my backyard, that had grown three tails, or partial tails. I have an update. Not only is he still alive, he now has a fourth piece growing. I noticed he had an injury to his tail about a month or so ago, and now a new piece is growing out of that spot. i remember a lot of people asked to permission to share the last photos i took. Please feel free to share. Taken in Broward County, FL, with a Nikon D7000 and a Tamron 90mm SP Di lens.”

Anybody ever seen anything like this?


Great Lizard-Watching Binos on Sale


My favorite brand of binos for lizard watching are steeply discounted right now. In optics–binoculars, cameras, etc.–you get what you pay for: more expensive units are generally of higher quality. Nevertheless, there’s a sweet spot that maximizes your bang for the buck, and that sweet spot in binos, I contend, is Eagle optics 8×32. Normally retailing at $380 or so, these have high quality lens and a good feel in the hand. And, most importantly, they can focus on lizards three feet away! Many binoculars are made for bird watchers and can’t focus anywhere near that close, making them not useful for looking at small objects such as lizards. And now the good news: the glasses are currently discounted to $239. Get ‘em while the supplies last! You might also consider the 10×32’s. They don’t focus as close–a respectable 8′ according to the specs–but they do give greater magnification. And they’re more than half off.