Author: Simon Lailvaux

Eye of the /Tiger Green Anole

In that classic of American cinema, Rocky III, Rocky Balboa (Sylvester Stallone) employed a particularly cunning strategy during the climactic fight with the younger, stronger Clubber Lang (Mr. T): he used his face to repeatedly absorb all of Clubber’s most powerful blows until Clubber grew very tired. Rocky’s strategy worked, and Clubber, fatigued from what seemed like hours of savagely beating Rocky in the head, ultimately succumbed to one of the relatively few punches Rocky managed to land.

Repeat until World Champion

Repeat until World Champion

Now one might suspect that Rocky III’s inspiring message of never giving up being punched in the face would have few adherents in the animal world, and this is indeed what we find. In most cases of male-male combat, combatants are reluctant to enter into escalated physical altercations because the risk of injury to themselves is too high. Instead, males of many animal species have evolved ritualized aggressive signals or displays aimed at intimidating their opponents into withdrawing, and will turn to violence only as a last resort when all else has failed. But some species have adopted the spirit of Rocky’s strategy, if not the letter, and rely on persistence to outlast as opposed to outfight their opponents.

A new study by Wilczynski et al. shows that Anolis carolinensis (the undisputed greatest study organism in the world) may use persistence as part of its fighting strategy as well. Adult male green anoles establish dominance hierarchies initially through aggressive interactions, and the outcomes of these interactions are affected by a variety of behavioural, physiological and morphological factors, many of which are likely reflected in the pattern and intensity of their ritualized aggressive displays. Wilczynski et al. set up staged aggressive interactions between pairs of adult males in the laboratory and tested whether males that responded faster or for longer to behavioural challenges were more likely to win fights. They also noted the colour state, as well as the presence of post-orbital eyespots, of winners and losers, both of which have been the subject of previous discussion on Anole Annals. figure 2The authors found that for the measured types of display, future dominant individuals generally displayed more frequently, and continued to display for longer than future subordinate individuals, whereas the effects of latency to display on competitive outcomes is less clear. With regard to colour, despite some intriguing trends, there were no significant differences between dominants and subordinates in any aspect of post-orbital eyespot expression. However, future dominant individuals did remain bright green for longer throughout the interactions than did future subordinates, supporting earlier suggestions that dark brown colouration is linked to subordinate social status and/or stress.

While persistence is a key component of contest behaviour in many animal species, the apparent importance of persistence in display duration in particular is especially interesting within the context of lizard displays. For example, duration of sagittal compression has previously been suggested as a handicap display in Uta stansburiana lizards, and previous studies have also suggested that persistence, perhaps related to accumulation of metabolic costs (paper here), might also dictate male contest outcomes in green anoles. Despite the wealth of knowledge regarding male green anole displays, studies such as Wilcynski et al.’s show that we still have much to learn regarding the behavioural aspects of male combat in this species, not to mention the likely relationships between behaviour and physiology.

Rocky III was unjustly spurned by the Academy of Motion Picture Arts and Sciences in 1983, not even receiving a nomination in the category of best picture (Ghandi won that year for some reason). Even more outrageous, it didn’t win the Best Original Song category it was nominated in! (Would anyone seriously argue that “Up Where We Belong” is a better song than “Eye of the Tiger”? Because it isn’t, and you are wrong). In retrospect, the reason for this travesty is clear: persistence is an important part of animal fighting strategies, and Rocky III was actually a nature documentary.

Are You Displaying At Me? There’s No One Else Here…

Three stereotyped head-bobbing patterns of A. carolinensis. The black area represents the amplitude of the head as it moves up and down. Although amplitude can vary within and among individuals, the cadence remains constant for each of the display types. The hatched area refers to times when the dewlap is displayed. Length and number of dewlap displays and associated headbobs at the end of the display can be quite variable. This figure is a modification from Jenssen et al. (2000) that appeared in Lizards in an Evolutionary Tree.

Anole displays have proven to be as frustrating as they are fascinating. These displays, which are species-specific and typically involve headbobs and/or pushups that may or may not be accompanied by dewlap extensions, are as varied as one might expect within a group as diverse as Anolis. However, despite concerted efforts by an energetic group of researchers aimed at understanding the form, context and meaning of these varied adult male displays, a complete understanding of this complex signal eludes us. The stereotyped displays exhibited by many anole species are of particular interest, and of these, arguably the best-studied are the A, B, and C displays of that lab rat of the anole world, Anolis carolinensis.

In a recent paper, Jenssen and collaborators report a characteristically detailed and rigorous field study on the use of such stereotyped displays in both breeding and postbreeding free-ranging A. carolinensis males. Through painstaking analysis of videotapes of thousands of undirected male displays in nature with no obvious receiver, they show, among other results, that breeding males in “monitor” mode (i.e. lizards that signal while stationary) used mostly C displays, but increased use of A and B displays while moving or “travelling.” Monitoring lizards also exhibited an overall lower display rate than travelling males, and used more and longer volleys of displays. Additionally, about a third of all displays had an extra “shudderbob” tacked on. These patterns held through the postbreeding season. Interestingly, Jenssen et al. note that these undirected displays aimed at no-one in particular are most similar to aggressive signals used by males engaged in contests. The implication is that these undirected displays are in fact directed at an unidentified (or undetected) rival male audience, rather than being for the benefit of any single lady lizards in the area.

The notion of males displaying aggressively just in case any rivals might be present makes sense for a species such as A. carolinensis that defends areas harbouring females, rather than trying to attract them. One wonders if this result would hold for species that place less emphasis on territory defense and that have been rumoured in the past to exhibit signs of female preferences (Anolis valencienni, anyone?).

Sizing Up Green Anole Dewlaps

Several years ago I was involved in a study showing that the dewlaps of individual male green anoles change size over the course of a breeding season, increasing in area from winter to spring and then shrinking from spring to winter. This result was first noted in the field and verified in the lab, and is not a statistical artefact – individual dewlaps really do change size!

Shortly after that study appeared I found myself in Australia doing postdoc work on crickets. During that time I gained an appreciation for life-history and the battery of approaches, ranging from artificial diets to mating schedule manipulations, which researchers use to expose resource allocation priorities in animals. (On a related note, I also gained an allergy to crickets). When I returned to the lizard world I started thinking about dewlaps and resource allocation, and I wondered if it might be possible to apply some of these life-history techniques to anoles to figure out the mechanisms underlying the incredible growing/shrinking dewlaps.

It turns out that not only is it possible, it’s actually pretty easy, and my research group was recently able to conduct a simple dietary restriction experiment that yielded some unexpected results. We wanted to test whether dewlap size is affected by resource availability,

The Perils of Urban Anolology

One of the perks of working on anoles is doing fieldwork in exotic and interesting places. For me, being located as I am in steamy New Orleans, an additional perk is that I can (and do!) do fieldwork in my metaphorical back yard. As of last year, I have begun what I hope will become a long-term study of a local population of Anolis carolinensis in Washington Square Park in the Faubourg Marigny, right on the edge of the French Quarter. (As an aside to fellow anole biologists, if anyone is planning future anole collecting trips to NOLA my lab and I would be more than happy to help out, with the caveat that the anoles in Washington Square Park remain uncollected and unmolested, given that selection is one of the many things we are currently measuring). But doing fieldwork in populated urban areas also presents its own unique set of challenges, not the least of which is the colorful local populace. This generally breaks down into several categories. In most cases, people are simply curious as to why someone would spend a warm, spring day diving into bushes for no apparent reason, and immediately assume that you are a lunatic once you explain to them that it is with the aim of documenting the Lizard Invasion (“They’re all around us!”). Others are genuinely fascinated with the anoles, and will listen with interest to what we are doing and why before leaving us to go about our business. Then there is another small segment of the population who hold strong and invariably uninformed opinions on science, lizards, nature and animal rights which they eagerly thrust upon hapless anole investigators with no warning or provocation. Dealing with these people can be difficult, as it requires certain baseline levels of patience and diplomacy that I sometimes do not possess. Finally, New Orleans being New Orleans  also plays host to a surprisingly large amount of people who are not burdened with an abundance of sobriety at 9am on a Tuesday (or, perhaps, ever). Interactions with these special people run the gamut from mildly amusing (the Grateful Dead fan last week who casually mentioned that he was on day 3 of “tripping [his] balls off” and helpfully warned me to “watch out for roofies”) to tiresome (the heroically inebriated gentleman who followed me around the park recently for 10 minutes whilst repeatedly slurring the same unintelligible question in my general direction) and even vaguely unnerving (the cracked-out, half-naked couple chasing a rubbish truck down Frenchmen street and yelling at it).

Pictured: Outreach

It was this inconvenient presence of the public in public places that in 2004 led Duncan Irschick, weary of interacting with other human beings, to make perhaps his most important contribution to anole biology – the “I’m catching lizards” t-shirt. Originally made for Irschick lab members during sampling of the Tulane University A. carolinensis population, these three simple words, printed in stark white on the back of a dark green shirt, were intended to inform the casual passersby as to the nature of our unusual outdoor activities, thereby dissuading them from engaging any of us in conversation. It works extremely well, except on those occasions when it backfires, instead acting as crazy bait and attracting exactly the type of person you don’t want to speak to whilst conducting fieldwork. Still, the benefits outweigh the occasional costs, and my shirt has seen regular use over the years. I don’t particularly recommend wearing it in non-lizard catching contexts, however, as herpetologists are generally considered to be weird enough as it is.

Powered by WordPress & Theme by Anders Norén