Tag Archives: Dewlap

Signals and Speciation: Do Dewlap Color Differences Predict Genetic Differences?

Dewlap and genetic differences between co-occurring Anolis distichus and A. brevirostris

Dewlap and genetic differences between Anolis distichus and A. brevirostris at sites where they co-occur on Hispaniola.

Here at Anole Annals, we’re all familiar with the replicated evolution of different anole ecomorph types in the Greater Antilles. However, divergence into these different ecomorph classes is not enough to explain how the group became so speciose on these islands. Additional factors must therefore have promoted speciation throughout the history of the group.

One potential factor is the flashy anole dewlap. Dewlap diversification across anoles has led to the remarkable array of dewlap color, pattern and size we see today. If dewlap differences did indeed drive speciation in anoles, or are involved with the maintenance of species boundaries, we might expect that as differences in dewlap color and pattern increases between species, genetic differentiation will also increase through fewer hybridization events.

In our study that just came out in the Journal of Herpetology, Rich Glor, Anthony Geneva, Sabina Noll and I set out to test this using two widespread species from the Anolis distichus species complex, A. distichus and A. brevirostris. These two species co-occur in many locations on Hispaniola and, while they often differ in dewlap color where they do co-occur (yellow with an orange patch vs. all pale yellow), in other areas, they co-occur with similarly pale dewlaps. Using mitochondrial DNA, microsatellite and AFLP data, we investigated patterns of genetic differentiation at four sites: two where the species differ in dewlap color, one where the species share the same dewlap color, and another where pale dewlapped A. brevirostris co-occurs with two A. distichus subspecies (one with a similarly pale dewlap and the other with an orange dewlap).

In general, we found that A. distichus and A. brevirostris looked like “good species,” with strong genetic differentiation and little evidence of hybridization, even at a site where they share the same dewlap color. This suggests that dewlap color differences are not associated with genetic differentiation in a manner one might expect if dewlaps were involved in the speciation process or in maintaining species boundaries. However, at the site where A. brevirostris co-occurs with two A. distichus subspecies with both similar and dissimilar dewlap colors, we found some evidence of hybridization and the species were not as highly genetically differentiated. This discrepancy suggests that site-specific factors could be influencing the dewlap’s role in speciation or maintaining species boundaries. For example, as Leo Fleishman’s and Manuel Leal’s work has shown (e.g. 1, 23), the dewlap’s effectiveness as a signal is dependent on the light environment. Further understanding about the environmental differences among our study sites, how species utilize the available light microhabitats within each site, and how the dewlap looks to anoles at each site could provide more insight into our findings.

On the other hand, perhaps we need to be looking beyond the dewlap and focusing instead on whole signaling displays. Anole behavioral displays can also be strikingly different among species (e.g. 1) and may instead be the key to understanding species diversification in Greater Antillean anoles.

Dewlap Displays in Cuban Knight Anoles (A. equestris)

While exploring the grounds of Fairchild Tropical Botanical Gardens with Janson Jones this past weekend, we extremely fortunately happened upon a large adult male Cuban knight anole (A. equestris) in full displaying swing. Despite the fact that knight anoles have an impressively large dewlap, I have often found this to be a relatively rare event, as large crown-giant species tend to display less than other smaller and more active species. This individual was displaying at a height of ~15 m, just below the fronds of a large Royal Palm (Roystonea regia). We didn’t see any other neighboring knight anoles, so were unsure if this was a directed or passive display series. In all, this lizard performed perhaps 4-5 sets of dewlap displays (each comprising of 4-5 dewlap extensions) before stopping and retreating back into the canopy.

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Anoles typically follow a predictable and repeated pattern of display that gradually increases in intensity. Initially, and rather lethargically, an individual will nonchalantly raise its head and extend its dewlap without much extra effort (stage a); described below from Losos (2009).

Adapted from Losos (2009), which itself is adapted from Losos (1985). Aggressive behavior of A. marconoi showing three stages of increasing display intensity - note stage (c) include full body elevation alongside simultaneous tail and dewlap extensions.

Adapted from Losos (2009), which itself is adapted from Losos (1985). Aggressive behavior of A. marconoi showing three stages of increasing display intensity – note stage (c) include full body elevation alongside simultaneous tail and dewlap extensions.

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This then escalates to include a slight body raise (stage b).

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And ultimately results in a dramatic finale – in full display all limbs will be extended to raise both their body from the substrate (in this case the trunk of a palm tree) and elevate their tail (stage c). In the following picture you can see this final stage of displaying where intensity peaks – albeit in this individual with a regenerated (and rather stubby) tail. Continue reading Dewlap Displays in Cuban Knight Anoles (A. equestris)

On the importance of Dorsal and Tail Crest Illumination in Anolis Signals

With a flurry of recent attention investigating how background light may influence the signalling efficiency of Anolis dewlaps (1,2,3,4), particularly those inhabiting low-light environments where patches of sunlight appear at a premium, it occurred to me that extended dorsal and tail crests may fall under similar selection. Below are some photos of Puerto Rican crested anoles (Anolis cristatellus) – a species in which males exhibit an enlarged tail crest and the ability to voluntarily erect impressive nuchal and dorsal crests during aggressive interactions (the mechanisms of which are detailed in this previous AA post) – that show how crests may contribute to signalling.

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I have no doubt this thought has crossed the minds of many anole scientists before, particularly those current graduate students so successfully studying A. cristatellus and familiar with their ecology and behaviour (namely Alex Gunderson, Kristin Winchell, Matt McElroy, and Luisa Otero). Dewlaps are undoubtedly of primary importance to anole signalling and communication, but what are people’s general thoughts on the relative importance of other morphological features?

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What’s All the Fuss About Dewlaps?

Anolis carolinensis from http://www.mascotissimo.com/wp-content/uploads/2008/02/anolis_carolinensis.jpg

A few years ago, Richard Tokarz and colleagues conducted a series of studies in which he surgically disabled the dewlaps of some male A. sagrei and discovered that these functionally dewlapless lizards had no trouble holding a territory and seducing females. In a new study, Henningsen and Irschick found that surgically reducing the size of dewlaps in male A. carolinensis by about one-third had no effect on male-male aggressive interactions in the lab. Makes one wonder what’s the big deal about having a dewlap.

Dewlap Color, Gene Flow, Habitat Specialization, and Speciation: A Tale of Two Contact Zones

Dewlap variation in Anolis distichus in Hispaniola. The photos at the bottom show the change in dewlap color along the two transects in the recent study by Ng and Glor.

Despite all of the research on anole evolution conducted in the last 40 years, one important question still eludes us: how does speciation in anoles occur? This, of course, is of fundamental importance, because the great species richness of these lizards implies that speciation has run rampant in this group. So, we’d like to know why.

We don’t know much about speciation in anoles, but we do know a little. First, it is thought that the dewlap plays an important role. Sympatric anole species almost never have identical dewlaps, and experimental and observational evidence suggests that anoles use their dewlaps for species-recognition. Hence, understanding anole speciation may, to a significant extent, reduce to understanding the factors that cause populations to evolve differences in their dewlaps.

A different perspective on anole speciation relates to the classic question of whether allopatry is necessary or whether, as suggested by many recent studies, natural selection driving differentiation—whether in allopatry or not—is a more important stimulus to genetic differentiation. Recent work in the Lesser Antilles by Thorpe and colleagues has argued that environmental differences are the primary drivers of genetic differentiation within anoles, a result also suggested by Leal and Fleishman’s studies on A. cristatellus in Puerto Rico.

In this light, perhaps the most enigmatic anole is Anolis distichus of Hispaniola. Continue reading Dewlap Color, Gene Flow, Habitat Specialization, and Speciation: A Tale of Two Contact Zones

How Many Times Have Lizard Dewlaps Evolved?

Polychrus gutturosus flashing its stuff. Photo from http://www.bijagual.org/images_reptiles/reptiles_image_links/pages/polychrus_guttorosus_3_JPG.htm

One interesting implication of the recent finding that Anolis and Polychrus are not closely related concerns the evolution of the dewlap. The two genera were long thought to be close relatives in part because they both possess what appear to be similar dewlaps. The new phylogeny indicates that these structures are not indicative of common ancestry, but rather that the two clades have convergently evolved very similar structures. 

Dewlap-like structures have, in fact, evolved repeatedly in iguanian lizards (the clade that contains iguanids [in the old, broad sense], agamids, and chameleons). Some of these dewlaps are different from that of anoles—such as the flap of iguanas and the triangular dewlap of Draco—but the dewlaps of the agamid genera Sitana and Otocryptis are dead ringers for those of anoles. In fact, one might argue that Sitana out-anoles anoles with its regal fan pictured below.

Sitana ponticeriana. Photo by Niranjan Sant from Lizards in an Evolutionary Tree

Continue reading How Many Times Have Lizard Dewlaps Evolved?

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?

Name That Anole (x2)!

I’ve enjoyed this type of post and figured I would contribute myself. On a trip to Costa Rica in early 2010, I had the pleasure of wandering around catching all the anoles I could see. Although most of my photos have unfortunately been lost in a massive computer/hard drive failure, I have recovered a few shots from the field. Here are photos (of two species) that always get me thinking about dewlap coloration, and maybe they will get you thinking more about that too. So what are the species, everyone?