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Why Are Some Brown Anoles Orange? A Laboratory Study

An orange Anolis sagrei used in the study. Image by Beth Reinke.

Readers of Anole Annals know that Florida populations of Anolis sagrei now include red-orange individuals [1, 2, 3]. I learned more about this new color by conducting the first scientific study on orange skin coloration in Anolis sagrei.

Before I go any further, I owe a thank you to those who documented their orange A. sagrei findings on Anole Annals. Previous posts confirmed that what I was seeing in the lab wasn’t an anomaly. As I learned more about the sightings of these orange anoles, it became apparent that the orange phenotype was rather common. The posts also helped me understand when this odd coloration was first noticed (only in the last decade!). I was even able to meet with one contributor in person.

The first thing I noticed was that there is quite a bit of diversity in the distribution of orange coloration on the bodies of the lizards themselves. Most of the posts on Anole Annals showcase full-bodied orange lizards [1, 2, 3]. I found that partial orange coloration was just as common. Take, for example, this male whose orange coloration was limited to his tail and hind legs.

A biologist’s first intuition is to wonder how differences in coloration might influence survival. Most of my research project was focused on identifying fitness differences between brown and orange lizards. I was working under the impression that orange skin suddenly appeared in the population and became common very quickly. I knew that there are cases when new phenotypes become common for no reason (genetic drift). Nonetheless, we don’t normally expect to see a new phenotype become common in a short amount of time. I suspected that orange lizards had an easier time surviving or breeding than the brown ones. But I was surprised that a color as conspicuous as orange could be so successful. I reasoned that it couldn’t have helped them camouflage, so why are orange lizards surviving and reproducing?

Maybe it had something to do with mate choice. Since males use their orange dewlaps to attract females, it might be that a completely orange male would look particularly stunning to a female. Even though orange might have made the males an easier target for predators, the effect on reproductive success may have outweighed the risk of predation. This is the hypothesis that I had in mind for most of the project and the one that made the most sense to me. It’s fitting, then, that when I ran a behavioral experiment in the lab, the females didn’t care at all about color! They were much more interested in males that performed a lot of pushups and head bobs (behaviors that many species of lizards use to communicate). These pushups and head bobs demonstrate a male’s physical fitness to a female.

Maybe orange reflected something in their physiology, then? I ran two different experiments to test endurance and sprint speed. The tests of endurance and sprint speed in particular took up most of the time of the project; it turns out live animals don’t usually do what you need them to do. Despite their penchant for sprinting out of sight in the wild, getting lizards to run in the lab was more difficult than you might guess. The endurance tests involved a custom-built lizard-sized treadmill. More often than not, the lizards would treat it like a moving sidewalk you’d find at the airport. Other times they’d wriggle into the machine itself (at no risk to them) and I’d have to take apart the treadmill, one screw at a time, to fish them out. No images of that, sadly.

To measure sprint speed, I needed the lizards to run up a wooden pole. Here’s a video of me trying to convince lizards to run up that pole.

I became more interested in paleontology after this project. Dead animals behave more predictably.

After all that, the data didn’t point to any difference in orange and brown lizards’ endurance or sprinting ability. I took a step back to get to the bottom of something I knew I could answer. I wanted to identify the pigments that they were using to color their skin. Having read about what gives Anolis sagrei dewlaps their red and orange color, I was expecting to see two classes of pigments in orange lizard skin: carotenoids and pterins. No one had extracted pigments from even brown A. sagrei skin before, but I wasn’t expecting to see much in non-orange skin.

I boiled lizard skin in all sorts of carcinogenic solutions to extract the pigments.
Then I separated the two types of pigments in test tubes – carotenoids at the top and pterins at the bottom.

As expected, the dewlaps had both types of pigments. Unexpectedly, brown lizard skin contained pterins. I thought this was a little odd since we don’t see red or orange on brown lizards. But, no one had done this before, so I didn’t quite know what to expect. Like brown lizard skin, orange lizard skin had pterins, but not carotenoids. This surprised me because it suggested that the orange color in orange lizards might not be due to the addition of a pigment so much as the absence of one. Melanin (another class of pigment that produces brown and black colors) typically masks the effects of other pigments that may be present. So, although I was unable to test this myself, I now suspect that the orange color is caused by a lack of melanin.

It was time to revisit that camouflage idea. I had taken for granted that orange was too conspicuous to conceal a lizard, but I needed the data to back up my claim. I collected quantitative data on brown and orange lizards’ skin color by using a spectrophotometer, which records color as the wavelengths of light reflected off a surface. The result is something that looks like this:

What A. sagrei dewlaps look like to a spectrophotometer.

One of my collaborators, Dr. Beth Reinke, applied these data to a visual model to predict how A. sagrei’s bird predators would see the new color. She identified that orange anoles are less conspicuous to bird predators. Now the strongest lead is what I had ruled out when I first began the project: camouflage!

So what’s up with orange A. sagrei? The color doesn’t make them more attractive to mates nor does it correlate to increased physical fitness. Because orange and brown skin contain the same kind of orange-producing pigment, my best guess for the mechanism is a lack of melanin in the areas that appear orange. And, although the new color looks conspicuous to humans, it may help orange individuals hide from bird predators. The benefits of orange as camouflage may explain why the new color persists in south-Floridian populations of A. sagrei.

There’s a lot left to know about orange anoles. A good next step would be to test the “orange as camouflage” result in the field. Additionally, research into the genetic basis of this phenotype may identify how it arose and the mechanism behind it. Some breeders have suggested that orange coloration is genetically dominant over brown coloration. This is something I wanted to identify in breeding experiments, but time ran out before I graduated from college.

Orange A. sagrei remain enigmatic. I hope to hear more about orange anoles from enthusiasts in the lab and the field!

Paper: Erritouni YR, Reinke BA, Calsbeek R (2018) A novel body coloration phenotype in Anolis sagrei: Implications for physiology, fitness, and predation. PLoS ONE 13(12): e0209261. https://doi.org/10.1371/journal.pone.0209261

Brown Anole (A. sagrei) Surveys in Orange County, CA

Louis Shanghan of the LA Times reports on Greg Pauly‘s field surveys of non-native Cuban brown anoles (Anolis sagrei) and geckos in Orange County neighborhoods

sagrei_california

“The anoles, which are native to Cuba, arrived here about a decade ago as stowaways in nursery plants,” Pauly said as the team strode down a leafy street, methodically scanning sidewalks, brick walls and tree trunks for the stick-like shapes of lizards basking in the sun. “Today, there’s at least 10 to 20 per residential lot in this neighborhood alone.”

“There’s a nice one over there,” he said, nodding toward an anole – about five inches long, adorned with light brown speckles and a bright line running from head to tail – clinging to the side of a front-yard planter box.

Full story here: Scientists survey an Orange County neighborhood’s nonnative lizard populations

As a side note, the details for the original record (as far as I know) of A. sagrei in California are as follows:

The first published documentation was in Herpetological Review 45(4), 2014, an edited version of which you can read below:

ANOLIS SAGREI (Cuban Brown Anole). USA: CALIFORNIA: San Diego Co.: Vista, elev. 158 m) 19 July 2014.
C. Mahrdt, E. Ervin, and L. Geiger. Verified by Bradford D. Hollingsworth. San Diego Natural History Museum (SDSNH 76128–76133).

New county and state record (Granatosky and Krysko 2013. IRCF Rept. Amphib. 20[4]:190–191)
Four adult males and two hatchling specimens were collected on a one-acre parcel landscaped with palms, cycads, and several species of tropical plants and ground cover. Several boulders scattered throughout the parcel were used as perch sites for male lizards. An additional 16 adults and six hatchlings were observed in the two-hour site visit (1030–1230 h). Adults were also observed beyond the property indicating that this population is established and likely expanding through the contiguous tropical landscaping of neighboring properties. According to the property owner, he first observed the species in August 2012 shortly after receiving shipments of palm trees in May–August originating from suppliers located in the Hawaiian Islands.

CLARK R. MAHRDT, Department of Herpetology, San Diego Natural History Museum, San Diego, California 92102, USA (e-mail: leopardlizard@ cox.net);
EDWARD L. ERVIN, Merkel & Associates, Inc., 5434 Ruffin Road, San Diego, California 92123, USA;
GARY NAFIS, (www.californiaherps.com).

More information on A. sagrei in California can be found here

Battling Crested Anoles (A. cristatellus) in South Miami, FL

While out watching lizards last week with my undergraduate research assistant extraordinaire, Oliver Ljustina, and fellow SoFlo anole Ph.D. student Winter Beckles, we happened upon a pair of male crested anoles (Anolis cristatellus) ready to rumble! This is quite early – but not unheard of – in the season for the commencement of territorial disputes, so it was a surprise to see them locking horns so aggressively. This couple were battling fairly high in the tree, at approximately 3m.

Anyway, here are the pictures!

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Anolis sagrei Survey Continued: Eleuthera, The Bahamas

beach scrub and bay scenic 2

I just got back from a short trip down to Eleuthera in The Bahamas where I was assisting Anthony Geneva (Harvard post-doc) in sampling lizards. Also along for the trip were Sofia Prado-Irwin (Harvard Ph.D. student) and Rich Glor (University of Kansas). We went with the main goal of sampling Anolis sagrei from four habitat types found commonly in the Bahamas as an extension of an ongoing project in the Losos lab (previous posts from: Rum CayConcepcion IslandRagged IslandBiminiMangrove habitat, and Great Isaac Cay). Specifically, we were looking to sample Anolis sagrei in mangrove, secondary coppice forest, closed coppice forest, and beach scrub habitats. These habitats differ in the height of the canopy, density of the understory, and composition of plants.

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We focused entirely on the southern half of the island near Rock Sound and Cape Eleuthera. We were successful in sampling two beach scrub habitats, two mature coppice forest, one secondary coppice forest, and one mangrove habitat. We were able to catch all four of the anole species found on Eleuthera: Anolis angusticeps, Anolis distichus, Anolis sagrei, and Anolis smaragdinus. We also encountered a number of other native herp species: the Bahamian boa (Chilobothrus striatus), Ameiva auberi, Eleutherodactylus rogersi, curly tailed lizards (Leiocephalus carinatus), and the Bahamian racer (Alsophis voodoo), as well as a couple of non-native species: Cuban tree frog (Osteopilus septentrionalis), and Hemidactylus mabouia.

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In my own research I work with Anolis cristatellus, the Puerto Rican crested anole. I am always surprised when I catch A. sagrei by how much smaller they are than A. cristatellus, although very similar in appearance otherwise. On this trip, I was also surprised that the A. sagrei, as well as the A. angusticeps and the A. smaragdinus, appeared to be much smaller than those I had encountered on Bimini last spring.

We also found that the density of lizards was quite low compared to what we expected and what I had experienced in Bimini, both during the day and at night. In all four of the habitat types, we saw an abundance of hatchlings, juveniles, females, and small males, but relatively few full adult male A. sagrei. For A. angusticeps and A. smaragdinus, we encountered only a few individuals total during the week of sampling. This reminded me of an odd experience I had last fall in Puerto Rico with A. cristatellus. It was the same time of year and I had an extremely difficult time locating mature animals in sites where I had previously sampled large numbers during the spring and summer months. Instead, I observed a large number of very young animals and females. I’m curious if this is a coincidence or if perhaps there is a strong seasonal effect on either male behavior (i.e., reduced visibility outside of the mating season) or male abundance (i.e., reduced numbers because of mortality during the mating season). In other words, are the males still there, but hiding, or are they really lower in abundance in the late fall? Or maybe I was coincidentally unlucky on both trips… I am very curious to hear thoughts on this!

Anolis sagrei using coral ground habitat.

Anolis sagrei using coral ground habitat.

Finally, I want to end with a short natural history note on the habitat use of the A. sagrei in the mangrove habitat. In this habitat we observed A. sagrei using perches at drastically different heights: some were 6 feet up, others were on the ground. Interestingly, the ones on the ground did not appear to be in transit, but seemed to be using the pockmarked karst as perches, running into one of the many holes when approached. Has any one else observed this behavior before? It seems so different from the typical trunk-ground anole perch and behavior to me.

That’s all for now. Currently Anthony is sampling additional islands in the Bahamas along with Melissa Kemp (Harvard post-doc) and Colin Donihue (Yale Ph.D. candidate / Harvard visiting student). Best of luck to them, I can’t wait to hear how the rest of the trip went!

Nephila Predation on Brown Anole

A brown anole is caught up in the web of an Argiope orb-weaving spider

A brown anole is caught up in the web of an Nephila orb-weaving spider

Anoles eating spiders and spiders turning the table on anoles are well reported in both the literature and here on Anole Annals (1, 2). Recently, biologists Sarah French and Matthew Wolak of UC Riverside encountered this unfortunate Anolis sagrei that had been caught up in the web of an Nephila orb-weaving spider. Here’s what they had to say about the enounter: “We were at the Gumbo Limbo Nature Center in Boca Raton. Matt & I were walking down the boardwalk, totally creeped out by the abundance of spiders, when we encountered the anole caught in a web. He was still alive, but pretty well caught. The spider didn’t seem entirely sure what to do with it, but she seemed to occasionally bite it, which caused the anole to jerk & thrash about for a few seconds. It was hard not to feel sympathy for the anole, but native species trump exotic, and so we refrained from interfering! (But we also didn’t stick around for too long to watch).”

A Morning Of Territorial Confrontations

As I photographed an A. carolinensis displaying high on a tree trunk, an A. sagrei popped out about 5 feet below and countered with a display. Before he could advance on the green anole male above, another male A. sagrei advanced to challenge. The two A. sagrei got in each other’s faces, but did not actually lock in combat. Suddenly the first A. sagrei broke off and advanced up the tree to confront the male green anole. There was a lot of counter displaying but not as fierce as just performed by the two brown anoles. Eventually the green male retreated further up the trunk, stopped to display once before disappearing around the other side.

Species-Specific Feeding Behaviors!

As an evolutionary biomechanist that is half in the Losos lab, I naturally dabbled in studying anoles during my first semester. I never presented my research, and have since moved on to other animals, but I thought you might like to see what I found.

Thom’s work on head shape shows a great amount of variation in the jaw length and width among anoles, and we wondered if the shape had an effect on jaw function. I was looking for differences in feeding behavior between the short-snouted Anolis sagrei and the pointy-snouted Anolis carolinensis. I placed a cricket on a leash, put it on a wooden perch inside a plexiglas container, put the lizard on the perch at the other end, and filmed the result.

Here are some videos of one sagrei attack:
sagrei- Front view

sagrei- Side view

And here is a video of a carolinensis:
carolinensis- Side view

Based on my limited dataset, it looks like the sagrei keep their heads low on the perch while they make an attack-dash consisting of 1 chomp. They hold the prey in their mouths for a while before they begin chewing. Carolinensis get very close to the prey, pause, raise their heads up, and stab their jaws downwards without moving their hind legs.

By the way, if you need ideas on how to study anole biomechanics, I’d love to chat!

Orange Anolis in South Florida

New minor color variants appear every once in a while, but it’s always interesting to find something completely different.  This, to the best of my knowledge, is something completely different.  I’ve found a few of these guys running around, and most had very similar colors.  Considering their size (and presumptive age) I wonder if they were from the same clutch, or if a single breeding pair yielded this Punnett square anomaly.

I think the concept of cryptic coloration isn't on his mind.

Both of the males I had time to annoy/photograph (and the one female that was slightly less photogenic) exhibited the usual traits of A. sagrei.  From the heavier build and shorter snouts, as well as the bolder attitude than our native carolinensis (I think the dewlap display was more for me than anything else; even when I was three feet away with a rather bulky camera, both males stood their ground), they would definitely fit the profile. But they’re not structurally an exact match to sagrei’s either. I don’t have a great head-on shot, but they’re narrower.  Considering the insect population in the area I can’t say it’s from undernourishment.  They move and jump more like carolinensis as well. They just don’t seem to be a differently-colored sagrei.  Maybe there’s a little A. cristatellus in there.

So what exactly is our bold little friend here?

I’m not the first one here to wonder what hybridization would yield and what cool little recessive traits could come from it, but I haven’t seen nearly enough specimens to suggest it’s a morph that may stick around- whatever it’s source.

Unique dewlap?

I recently returned from a trip to eastern Cuba and as expected, made some interesting observations and gathered some new natural history information.
While poking around one evening with a flashlight (mainly looking for Eluth’s) I saw this “orange” sagrei sleeping on some veg. I photographed it to share here since there was some discussion on and off blog about this color phase. After I got it in hand to determine species (since homolechis and jubar were also very common in the area), I was surprised at the dewlap appearance. At first I thought it had a red mite infection because of the color and texture; but after scrutiny, just accepted that it had a bright red pigment that was scattered about the entire ventral anterior. Any ideas or similar observation?

Anoles in the Blogosphere

It turns out that Anole Annals isn’t the only member of the WordPress.com stable that has a thing for our favorite lizard. While recently doing some tag surfing, we came across the following posts.

Catholic mom tells the gripping (or not) story of a green anole that went for a ride on the minivan windshield. You can probably guess the outcome, but the photos are nifty.

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