Tag: Draco

Dewlap Displays Supersede Headbobs, Yet Again

On March 11, 2023

The dewlap is probably the most noticeable thing about anoles. For me, the best way to spot an anole is by the flash of color from the dewlap as a lizard displays. Without that, many anoles would remain cryptic amongst the vegetation. This seems to be the case for the lizards themselves as well. The burst of color and movement as the dewlap is rapidly extended is a wonderful device for attracting the attention of rivals and mates. It’s possible that the dewlap originally evolved as an attention-grabbing flag to augment an existing sequence of elaborate headbob movements in forested environments. These days, the dewlap is a complex signal component in its own right, often with a dizzying array of colours and displayed using a variety of movements.

Anoles aren’t the only ones with a moveable dewlap. The Southeast Asian Draco lizards have a dewlap, and again to back up the headbob movements that make up their main channel of social communication. There are many other parallels between Draco and Anolis lizards, but the similarities in how they communicate is something that particularly fascinates me.

Early on in my fieldwork with Draco, I started discovering species that didn’t seem to use headbobs as part of their social display. It seemed these species had lost the headbob entirely and instead concentrated all of their communication through the dewlap display. These species are a minority, but not by much. It was a puzzle. These Draco had lost a central and complex element of their communication in favour of something that was seemingly more basic. Communication biologists are often fixated on trying to explain how animal communication becomes more elaborate over evolutionary time, but less attentive to why complexity subsequently becomes lost. These Draco lizards were an excellent case study.

Draco melanopogon (photo above) only communicates using the dewlap, whereas Draco sumatranus (opening banner photo) relies on both headbobs and the dewlap, just like anoles.

After nearly a decade of fieldwork on numerous species of Draco throughout Malaysia, Borneo and the Philippines, my trips stalled in 2020, as did the rest of the world. Celebrities had nothing better to do than write biographies, but my lockdown project was to focus on using the data I already had at hand to finally solve the curious case of the missing headbob.

It felt like an endless series of stay-at-home orders in Australia, and well into 2021 too. While the celebrities had gone on to finish their books and were now doing the zoom promotion circuit, my progress was hurdled by home-schooling two young children. We survived home-schooling in the end, and my attempt at figuring out why some Draco have lost the headbob has finally been published.

The evolutionary history of visual displays in agamid lizards

The first discovery is the headbob display is very ancient, evolving something like 130 million years ago or more. That’s before the evolution of Draco, and before the evolution of the anoles, in an evolutionary ancestor to both the iguanid (new world) and agamid (old world) lizard families. This was back in the age of the dinosaurs. Today, virtually all iguanid and agamid lizards use a headbob display or some variant of it in social communication. Which means the absence of the headbob in a handful of Draco species is very unusual.

The loss of the headbob from the social display of Draco is effectively a loss of complexity. A loss of complexity means a loss of “information potential.” Try writing a biography with half the alphabet. You might manage the following or something a little longer: “I was born. I paid taxes.” Thirteen unique letters in total. Obviously not the rich backstory you might hope for. Not because you hadn’t lived a fulfilling existence; rather you don’t have the language complexity to convey it in detail.

There are various reasons animals might lose complexity in their social signals. Perhaps the original need for a complex signal is no longer present. Perhaps the invasion of a new environment puts a brake on the level of complexity that can be accurately perceived. Or perhaps natural selection on other things, like body size, has made performing a complex signal too costly.

The beauty of having spent so much time in the field is the accumulation of a large library of data. By leveraging this information, I was able to test each of the above scenarios. The short of it is, Draco that have lost the headbob are unusually large species. Physically moving the head and body in a headbob display is more energetically expensive than pumping the dewlap in and out. It seems, then, that the physiological cost of performing the headbob became too great for these large species and they shifted to relying only on the dewlap for communication. This implies the communication system of these species is compromised, unless they have made up the loss of information potential somewhere else.

Draco without the headbob have more complex dewlap colour patterns. Each dot is a different species.

In fact, the dewlap itself tends to be more complex in Draco that have lost the headbob. Stealing a method for measuring complexity of anole displays, the dewlap of these Draco are more elaborately coloured than the average Draco. Unfortunately, this is unlikely to have been enough to fully cover the loss of the headbob. This means Draco that no longer use the headbob are relying on a constrained communication system.

The idea that the headbob is likely to be more energetically expensive than the dewlap was originally proposed for the anoles. It was used to explain the physiological basis for why Jamaican anoles might have evolved an innovation that allowed them to move away from a headbob-centred display in favour of one focussed on the dewlap. To be clear, the Jamaican anoles do still rely on headbobs in their social displays. But a rapid series of dewlap pumps features more prominently in their displays compared to the typical anole, like those on Puerto Rico for example.

It seems the dewlap has begun to supersede the headbob in anoles as well.

If you’d rather not slog through the paper itself, you can view a 12 minute video summary instead. If you would like to slog through the paper and can’t access it behind the paywall, drop me an email and I’ll forward you a free copy (t.ord@unsw.edu.au).

Dewlap Size Is Not What We Thought

By Terry Ord

On October 14, 2022

In All Posts, New Research

The large, colourful dewlap is an obvious defining characteristic of the anole. Understandably, then, there has been a lot of investigation (and speculation) on what the dewlap is used for. Without doubt it’s for social communication, but to communicate what. Historically, the dewlap was thought to be used for species recognition, which remains a reasonable explanation today. But a typical assumption made by many anole researchers and evolutionary ecologists alike is the dewlap, and specifically its size, is effectively an ornament used to attract mates or advertise potential fighting ability among territorial rivals. In other words, the evolution of the dewlap is the product of sexual selection.

If that’s the case, then dewlap size should be linked to some aspect of an individual’s ‘quality’ or physical condition, especially in males who seem to be the ones courting females (not vice versa) or defending territories. This is because a male’s quality or condition can be hard to assess by general appearance alone, unless there is a key feature that provides an honest indicator of that quality. In anoles, this is assumed to be a large dewlap that’s physiologically costly to produce.

One easy way that has been proposed to test for sexual selection in the origin of a morphological structure like the dewlap, is to look how it scales with body size. Structures that are honest indicators of condition will be costly to develop and maintain. Large males are often in better condition than small males because of the underlying factors that result in bigger bodies (e.g., a history of successful foraging, superior growth rate, having ‘good’ genes). This means larger males can invest more in exaggerating the size of the dewlap than smaller males. There would be a clear evolutionary incentive to do so as well, because having a larger dewlap would attract more mates and appear more threatening to male rivals. The outcome of this should be disproportionately larger dewlaps in larger males. This is called positive allometry or hyper-allometry. If dewlap size has a hyper-allometric scaling relationship with body size, then it probably resulted from sexual selection. Or at least that’s the idea. And you can find this out by just measuring a bunch a males.

The dewlap of anoles featured heavily in the original formulation of this idea, with the conclusion being that dewlap size was hyper-allometric and assumed to be the product of sexual selection. Anoles have therefore become a classic example of how sexual selection drives hyper-allometric scaling in ornament size.

Tom Summers

Tom Summers was a graduate student who thought about hyper-allometric scaling a lot. He looked at the scaling relationship of ornaments that he had confirmed experimentally to be the target of sexual selection in fish, and found they were hyper-allometric…sometimes. Tom found natural selection on ornament size can often work in the opposite direction to sexual selection. This is because large ornaments can interfere with locomotion and often be conspicuous targets for predators. When these pressures are high, species tend not to show hyper-allometry in ornaments. Those ornaments were still the product of sexual selection, but their allometric scaling was dampened by opposing natural selection.

Tom turned this attention to the anoles, and found overwhelmingly that dewlap size was not hyper-allometric but hypo-allometric. That is, larger males have disproportionately smaller dewlaps than smaller males. He even looked at another group of lizards that have independently evolved a dewlap, the southeast Asian Draco, and found the same pattern. His results have just been published in the Journal of Evolutionary Biology.

The scaling relationship of the dewlap in both groups varied from one species to another, but never was it hyper-allometric. In the case of the anole dewlap, this variation in dewlap size was predicted by factors important in signal detection (receiver distance and habitat light). This was consistent with the general hypo-allometry of the dewlap as well.

The effectiveness of a visual flag (like the dewlap) in attracting the attention of a receiver (another lizard) is dependent on the gross size of that flag, not how big it is relative to the signaller’s body (i.e., allometric scaling is irrelevant). Beyond a particular threshold size, which is dependent on the visual acuity of the animal in question, there are diminishing returns for detection with increasing size. Even a large increase in dewlap size beyond a certain point wouldn’t really improve signal detection, a phenomenon known as ‘Weber’s Law’. The resulting pattern when comparing dewlap size among males is hypo-allometric scaling. Larger males have generally reached the size threshold for reliable detection, so there’s little point in further elaboration.

It also fits with the extensive amount of work showing that the dewlap is likely to be most important in signal detection, rather than a cue of quality.

So why such a dramatically different finding to earlier investigations of the anole dewlap? All studies prior to Tom’s measured dewlap size by catching the lizard and manually pulling out the dewlap using forceps. Simon Lailvaux has discovered that the skin of the dewlap varies in its elasticity. Larger dewlaps are going to be more stretchy than smaller dewlaps. This means you can probably pull the dewlap out to a larger size in larger males. This would subsequently generate the artifact of hyper-allometric scaling when comparing dewlap size across males of different size.

Tom had measured dewlap size from high-definition videos of free-ranging males fully extending their dewlaps during display. There are various analyses in his paper that confirm this approach provides an accurate measure of dewlap size. His logic at the time was this view of the dewlap would be how lizards actually see and evaluate the size of the dewlap relative to body size. It also meant animals didn’t have to be caught, so the approach was less intrusive for the animal (always a plus). It just happened he avoided the potential problem of over stretching the dewlap if he had caught the animals and manually extended the dewlap by hand.

What does this mean for all that data that has been based on researchers pulling out the dewlap using forceps to measure its size? Honestly, I don’t know. Maybe nothing depending on what the data are being used for. Maybe everything if the data are being used in allometry studies.