Tag: Dewlap Page 1 of 2

Dewlap Displays Supersede Headbobs, Yet Again

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).

#DidYouAnole – Anolis allogus


Photo: Alex Alfil, iNaturalist

Hello!

Happy Holidays and Happy Anole Eve!

I took a break for my birthday, and then it was a holiday, then finals, then I had to travel, but now I’m back and it’s almost a holiday again. Before another holiday I did wanna share a short anole post! I’ve also missed doing this and I’ve been thinking about what anole to do since I got to see the anole specimens at the London Natural History Museum.

I picked a festive little Cuban anole, Anolis allogus, also called the Spanish Flag anole or Bueycito anole (after a village where it can be found).

The males of this trunk-ground anole species have an SVL of ~60 mm, while females are about ~49 mm. Its distribution is sporadic but it’s found mostly in the east of Cuba. They follow the typical trunk-ground anole coloration of light or dark grey-brown tones, with striping on their tails, and marbling on their sides. Male Bueycito anoles have pale yellow dewlaps with either a center dot or lateral center stripes of bright red/red-orange. Females have smaller pale yellow dewlaps.


Photo: Alex Alfil, iNaturalist

Spanish flag anoles have been recorded predating on a species of Blindsnake. Check it out here!

I know this was short but I just wanted to say hi really quickly and Happy New Year!

#DidYouAnole – Anolis phyllorhinus

Adult male specimen of Anolis phyllorhinus MYERS & cARvALHO, 1945,... | Download Scientific Diagram
Photo: Moares & Werneck, 2019

I think we may have to move #DidYouAnole to Fridays since that seems to be the better for me post recently.

And speaking of this week’s post I remember mentioning that there were other anoles with little rostral appendages and that I hadn’t gotten back to them.
(A shame it took me so long because they really are great anoles)

Anolis phyllorhinus, or the Leaf-nosed anole, is endemic to central Amazonia in Brazil (where I believe they’re called Lagarto papa-vento in Portuguese) but they are an uncommon sighting. They’re a great shade of leaf green, with pale green-white undersides. Like with Anolis proboscis, these anoles’ appendages are also flexible and possibly used to display.

The eponymous leaf nose is only present in the males, with female Leaf-nosed (or Bat) anoles not even having any swelling or prominence of their noses. Female Leaf-nosed anoles also have a greatly reduced white dewlap, while the males have a larger one that is bright red on the front half and blue-green or white toward the neck.

The SVL of a Leaf-nosed anole is about 71-85 mm, excluding the proboscis which itself varies from 20-23 mm in measured specimens.

#DidYouAnole? – Anolis alvarezdeltoroi


Photo: Wouter Beukema, iNaturalist

So I’ve been reading a lot of anole papers, aside from the ones I normally read for fun (can’t believe I read papers for fun now), and I found an anole that’s pretty similar to two anoles I’ve looked at before but also still unusual.

Welcome back by the way. Nice to have to you here again.

This week is a third little cave anole, Anolis alvarezdeltoroi, or the Mexican cave anole. Mexican cave anoles live in a similar karst limestone habitat like Anolis bartschi and Anolis lucius and are often found deep inside caves, occasionally sleeping from the roof it. They may also perch from vegetation in or around the caves, particularly as juveniles.


Photo: Daniel Pineda Vera, iNaturalist

Like the other two anoles, the Mexican cave anole has a similar short body/long hindleg morphology. In a paper redescribing the species, the average SVL of the male anoles they measured was 53.3-74.0 mm, and 49.6-66.5 mm in females.

They seem to rely heavily on the karst habitat with healthy populations being found in areas with diminished forest but intact limestone/cave areas.

Male Mexican cave anoles have dark red dewlaps with white lateral rows of scales, while females have smaller black dewlaps with a similar pattern.


Photo: Arístides García Vinalay, iNaturalist

Please read the paper redescribing this anole here! For a while there was only a specimen available of it and not much info, but they worked on it and you should check it out. I wouldn’t have been able to write this if it wasn’t for them.

#DidYouAnole – Anolis rubribarbus


Photo: Alex Alfil, iNaturalist

I almost missed that it’s been a year of #DidYouAnole!! Thank you for sticking with me and learning about these lizards!

For this super belated one year anniversary, I have both an anole and a request.

First, the anole!

Anolis rubribarbus is a trunk-ground anole from Cuba. Specifically the eastern half of the Holguín province, around the town of Sagua de Tánamo from whence it gets one of its common names. The Sagua de Tánamo anole lives in the rainforest and possibly on rocky montane habitat in its range.

It’s listed as endangered due to its very limited range.

Also known as the Cuban tiger anole, their dewlaps are yellow with orange stripes and they are tan to olive with darker vertical stripes, like a… you know… tiger.


Photo: Alex Alfil, iNaturalist

Now for my request! I couldn’t find a lot about this anole’s ecology or natural history, so if you have worked on this species or know someone who does, please let me know!

In the meantime, here is a year in anoles from my Twitter, and from the blog!
Thank you! It means a lot to get to share these lizards with you.

SICB 2020: Sex-biased Parasitism and the Expression of a Sexual Signal in a Tropical Forest Lizard

Panamanian slender anole (Anolis apletophallus) (Photo Credit: Dr. Christian Cox)

Male sexual signals, and their often-associated distinct phenotypic and behavioral displays, have been hypothesized to have evolved from multiple sources. Two of which include the Good Genes hypothesis, which suggests sexual signals serve as an honest signal to potential mates, and the Immunocompetence Handicap hypothesis, which indicates trade-offs to elaborate signals. However, Dr. Christian Cox, an assistant professor in the Biology Department at Florida International University, thinks of these as a continuum rather than opposing hypotheses. The vibrant dewlaps of Anolis species serve as an excellent model system to address questions related to this continuum. Dr. Cox’s lab has documented sex-biased parasitism, which he discussed on Monday at SICB 2020. 

Dr. Christian Cox

The Panamanian slender anole (Anolis apletophallus) is known to host ectoparasitic trombiculid mites (also known as chigger mites). In this species, males and females are roughly the same size and, as with most anole species, males carry a large, colorful dewlap beneath their chins. Dr. Cox asked whether males and females of this species differ in the attachment site and intensity of mite infestations and whether any other factors (e.g., energetics) might influence infection. Using a combination of field and laboratory studies, Dr. Cox and his colleagues quantified the number of mites, prevalence, and intensity of infestation on individuals. He also measured mass, snout-vent length, and dewlap size. Following this, he collected fat bodies, livers, and gonads to investigate differences in energetics. 

Dr. Cox found that males were more likely to have mites on their dewlap whereas females had more mites in the inguinal and axillary regions. Additionally, large males had more mites than smaller ones and there was a significant correlation between the size of the dewlap and the number of mites. Dr. Cox also found that there was a negative relationship between fat body mass and the total number of mites. In other words, males with a heavier fat body had less mites. In females, there was a positive relationship between gonad size and the number of mites. These findings suggest sex-dependent factors influence ectoparasite load and are indicative of trade-offs to male sexual signals. 

Colour Change in the Gorgetal Scales of an Anole Dewlap

An adult male Anolis amplisquamosus with black gorgetal scales immediately after capture (left); the same individual ~10 min later with white gorgetal scales. Photo Credit – John David Curlis

 

Anole dewlaps are excellent examples of a “complex signalling system.” They exhibit a staggering diversity of colours and patterns. Each dewlap is species specific and adapted to enable these lizards to communicate, attract mates and guard their territories from rivals or competitors. Generally, the colour of a dewlap (and its gorgetal scales) is considered an unchangeable descriptive trait. This colouration is not only relied upon by scientists looking to identify a species, but also by anoles that co-occur and partition with different species in their select niche.

Therefore, it might be surprising to learn that recent observations prove rapid colour change in anole gorgetal scales is possible. The question is, what implications does this have?

A recent publication in IRCF Reptiles & Amphibians details an observation of Anolis amplisquamosus whereby a male individual upon capture possessed black gorgetal scales that quickly changed to pale yellow. Upon consulting the literature, it seems only one prior documentation of colour change in gorgetal scales was reported (Leenders and Watkins-Colwell, 2003), coincidentally also involving a member of the same species clade.

This recent observation of chromatophoric regulation in anole gorgetal scales may be significant in the wider context of anole biology, in confirming photographically that coloration is not always a fixed descriptive or diagnostic feature — at least among members of the A. crassulus species group. Accordingly, this information suggests that some anoles may have the ability to regulate the colour of their gorgetal scales in the same manner as they regulate dorsal and lateral scale colour.

Because the colour of gorgetal scales is a character often used in species identification, understanding the mechanics and the purpose of such a change is crucial; as well as any implications to display behaviour, communication and anole interactions.

SICB 2019: Do Bark Anoles Show Behavioral Syndromes?

Daisy Horr, an undergraduate researcher at Trinity University, discusses how bark anole behavior varies across several different social contexts.

Animals often use diverse behavioral repertoires to adjust to new, unexpected, or changing conditions very quickly. While it may seem like individuals could always use the best behavior for any given situation, we know that instead behaviors are often related within an individual. In other words, an individual’s behaviors are not always independent and may represent an underlying “behavioral syndrome” or correlated set of behavioral responses to related environmental conditions. These behavioral syndromes are also sometimes called “personalities” (though application of this word to animals can be a bit controversial!). So, for instance, an individual that has a “bold” behavioral syndrome might take little time to explore a new habitat or consume a novel food item more quickly, but also be more likely to stay active in the presence of a predator rather than hiding (the safer option!).

While anoles have been the focus of much behavioral research, we still lack an  understanding of the diverse behavioral phenotypes, including behavioral syndromes, which are displayed by a variety of anole species. The bark anole, Anolis distichus, is native to Hispaniola but also found in southern Florida where it has been introduced. While small, bark anoles can be quite feisty, and are known for their dramatic display behavior in the presence of male and female conspecifics.

Male bark anoles demonstrate pushup displays prior to engaging in combat.

Taking advantage of the bark anole’s willingness to put on a show, Daisy Horr, an undergraduate student and McNair Scholar in Michele Johnson’s lab at Trinity University, led a group of fellow researchers in assessing whether these anoles show behavioral syndromes. To do this, Daisy and her collaborators measured the degree to which male bark anole behaviors were repeatable across three different contexts: trials with another male present, trials with a female present, and solo trials in which no other anoles were present. They measured variables quantifying movement as well as display behaviors such as pushups and dewlap extensions during these trials. The team also wanted to see whether the measured behavioral traits were linked to morphological and physiological variables.

Daisy and colleagues found no support for the idea that behavior was linked to morphology, including size of the body, head, and dewlap, and mass of the whole body, the liver, and fat pads (structures holding fat as energy reserves), or the hematocrit of the lizards, a metric quantifying how many red cells are present in the blood. Bark anoles did show some level of behavioral consistency, however. Movement behaviors were quite repeatable even between trials with and without conspecifics. Display behaviors, however, including pushups and displays, were repeatable within, but not across contexts. This work suggests that bark anoles have consistent behavioral syndromes in some contexts. Looking forward, research into behavioral syndromes in anoles could offer insights into how behavior may vary with habitat use, ability to invade novel environments, or selection on behavior itself!

JMIH 2018: The Curious Case of Bark Anoles

The Bark Anole (Anolis distichus ignigularis) from the Río Recodo. Photo from Richard Glor’s Flickr.

The Bark Anole (Anolis distichus) is a highly polymorphic lizard widely distributed in Hispaniola. Anolis distichus is divided into 16 subspecies with dewlap colors ranging from deep wine red to pale yellow (Glor and Laport 2012). In the early days scientists, such as Albert Schwartz, argued that A. distichus is divisible into multiple subspecies according to an analysis of variation in body color and dewlap pigmentation. But, are they really subspecies?

During the 2018 Joint Meeting of Ichthyologists and Herpetologists (JMIH), Richard Glor shared his lab’s advances on the curious case of Bark Anoles. Anolis distichus populations have ecological, phenotypic and genetic differences. Previous studies show a correlation between dewlap phenotype and environmental variation; in drier habitats, lizards have smaller, brighter, yellow dewlaps, while those in wetter habitats have larger, less bright, orange dewlaps (Ng et al. 2012).

Previously, the Glor Lab found strong support for the hypothesis that A. distichus is comprised of numerous genomically distinct populations (MacGuigan et al. 2016). Genetic divergence was associated with a biogeographic barrier, but not with dewlap color. Also, they found evidence for hybridization in contact zones with limited gene flow and intrinsic reproductive isolation between subspecies (MacGuigan et al. 2016; Ng et al. 2016). Overall, these studies suggest that geographic isolation, as well as ecological specialization, contribute to speciation.

The Glor Lab continues putting together the pieces of the puzzle. Most recently, they sequenced and assembled whole genome sequence data for A. distichus to identify the genomic basis for species differences and speciation.

Drivers and Constraints of Within-Species Diversity in Dewlap Design

Sampling locations of the populations of study across the Caribbean. (1) Soroa (Cuba), population 1; (2) Soroa (Cuba) population 2; (3) Grand Cayman; (4) Santa Clara (Cuba); (5) South Bimini; (6) Chub Cay; (7) Andros; (8) Crooked Island; (9) Acklins; (10) San Salvador; (11) Staniel Cay; (12) Pidgeon Cay; (13) Grand Bahama; (14) South Abaco; (15) Cayman Brac; (16) Little Cayman; (17) Jamaica.

The dewlap is arguably one of most fascinating features of anoles. For me, it is the baffling diversity in dewlap size, coloration, and use —both among and within species— that makes it so interesting. However, understanding the origin and evolution of dewlap diversity in Anolis has proven a daunting task (Nicholson et al. 2007; Vanhooydonck et al. 2009). In an attempt to make (a little more) sense of the drivers and constraints of anole dewlap variation, a team of Belgian researchers from the University of Antwerp, led by evolutionary ecologist Tess Driessens, decided to look at dewlap diversity in Anolis sagrei. They surveyed 17 island populations of A. sagrei across the Caribbean and quantified dewlap design (color, size) and dewlap display behavior of both males and females.

Last year, Driessens and colleagues published their findings on how variation in abiotic factors (such as precipitation, temperature and other climatic variables) could explain much of the observed inter-island variation in dewlap design and use in A. sagrei (‘signal efficacy’ hypothesis). In a paper that came out last week, the team reports on the role of the biotic environment in driving dewlap diversity in the brown anole. Inspired by the wonderful study of Vanhooydonck et al. (2009), the researchers tested whether among-population dewlap variation could be (at least partially) assigned to variation in predation pressure (estimated by island size, tail break frequency, presence/absence of the predatory curly-tailed lizards, clay model attack rate), sexual selection (using sexual size dimorphism), and/or species recognition (number of syntopic Anolis species). Overall, they found only limited support for the idea that the extensive interpopulational variability in dewlap design and use in A. sagrei is mediated by variation in their biotic environment. Although they did find that males from larger islands show higher dewlap display intensities than males from smaller islands, and that males are more likely to have a ‘spotted’ dewlap pattern when co-occurring with a high number of syntopic Anolis species, the direct connection with predation pressure and species recognition remains ambiguous and demands further investigation.

In another recent paper, focusing only on the size of the male dewlap and their maximum bite capacity, the Belgian researchers asked a different question: does dewlap size signal fighting capacity (estimated by bite force) in A. sagrei, and is this true for all 17 sampled populations? And, does the level of signal honesty (that is, the steepness of the dewlap size-bite force relationship within a population) vary among populations, and is it linked with the strength of intrasexual selection? Their results showed that absolute dewlap size is an excellent predictor of bite force in all A. sagrei populations. However, relative dewlap size was only an honest signal of bite performance in 4 out of the 17 populations. Surprisingly, the level of signal honesty did not correlate with the strength of intrasexual selection.

Male brown anole biting on a purpose-built force plate. Photo by Tess Driessens

While the work of Tess Driessens and her team sheds new light on the drivers and constraints of dewlap diversity in A. sagrei, there is still plenty of study material left for future dewlap fanatics.

Page 1 of 2

Powered by WordPress & Theme by Anders Norén