I am an Assistant Professor of Biology at New York University. My first love is herpetology, but my lab works on urban evolutionary ecology in diverse organisms (but primarily anoles!).
If you missed the live Twitter broadcast of the final round, here it is: the conclusion of Anole March Madness 2016. In the final round, Anolis equestris represented the island anoles and Anolis frenatus represented the mainland anoles to answer the question every anole biologist ponders: are island anoles or mainland anoles superior?
Without any further ado, here’s the final match!
Our (fictional) anole tournament is nearing its end. The two semi-final matches concluded leaving only two anoles remaining. Who will be the ultimate champion, an island anole or a mainland anole?
Here’s the recap of the Final Four:
Winner match 25 (Anolis aquaticus) versus winner match 26 (Anolis frenatus)
In the rainforests of Columbia, Anolis aquaticus has ventured slightly out of his native range. Skittish from his previous streamside encounter with the giant Anolis insignis, he warily watches the canopy despite coming out on top in his last battle against Anolis limifrons. He is no stranger to Anolis frenatus, whose range extends north into Costa Rica, yet he fails to recognize the danger he is in as he climbs a tree towards a swarm of tasty insects. Out on a branch 10m up A. aquaticus gorges himself on the abundant food that tasted so much better than the streamside meals he was used to. Out of the corner of his eye, he sees the imposing form of Anolis frenatus move. More than twice his size, A. frenatus is a sight to behold. Anolis aquaticus quickly surveys his escape options. Seeing a stream below he rationalizes that he can escape if need be. He continues to gorge himself. Anolis frenatus is taken aback at the impertinence of this newcomer. He makes his way towards the branch upon which A. aquaticus is perched. Not willing to abandon his smorgasbord, A. aquaticus boldly stands his ground. He turns to face A. frenatus and begins to dewlap; perhaps this is a friendly giant that will want to avoid conflict. Anolis frenatus stops to size up this intruder and quickly determines that this tiny anole poses no threat to him. Anolis frenatus does not even bother to dewlap back. He continues on his path towards Anolis aquaticus, who stubbornly continues to dewlap and push-up in his cloud of tasty morsels. Anolis frenatus lunges at A. aquaticus, who instinctively turns and leaps without looking towards the stream. Anolis aquaticus sails through the air and lands safely in the water, or so he thinks. As he climbs onto a boulder, a fish-eating spider (Trechalea spp.) with a body size of nearly 17cm seizes the opportunity and leaps onto the unsuspecting and soggy anole. The spider makes quick work of the aquatic anole. Anolis frenatus watched from his high perch with a mouthful of delectable insects which he incidentally gobbled when he leapt at A. aquaticus with mouth agape. Anolis frenatus revels in his glory as he watches his competitor meet his end below. ***Anolis frenatus wins***
Winner match 27 (Anolis equestris) versus winner match 28 (Anolis bimaculatus)
A hurricane of unprecedented proportions has just ripped through the Caribbean. The Cuban Knight Anole, Anolis equestris, and the Panther Anole, Anolis bimaculatus, both find themselves washed up on an unfamiliar low-lying island. The waves periodically break over the island, washing away the debris on the ground. Both lizards quickly find a perch and scramble to escape an incoming wave. As the reach safety atop their scrubby perches, they catchy sight of each other. Clearly this island refuge has room for only one of them. Two lizards enter, one lizard leaves on this subsiding volcanic dome. They lock eyes and begin to dewlap. Nearly evenly matched it’s not clear who will claim this island as his new home. Anolis equestris extends his dewlap first. He unleashes a flurry of push-ups and rapid flashes of his cream colored dewlap. The slightly smaller Anolis bimaculatus has no choice but to stand his ground. He strains to extend his orange-yellow dewlap as far as he can. Unfortunately, his species has a disproportionately small dewlap for their size, and this A. bimaculatus is average sized at best. Anolis equestris, completely and utterly unimpressed, leaps from bush to bush with waves breaking over the rocky ground below him as he heads towards A. bimaculatus. Anolis bimaculatus makes an offensive move and lunges at A. equestris, who has by now reached the neighboring bush. Anolis bimaculatus lands a calculated blow on the abdomen of A. equestris. Despite the searing pain in his kidney, A. equestris reaches around and snatches A. bimaculatus in his forceful jaws. In one quick motion, A. equestris flings the slightly smaller A. bimaculatus over his shoulder and off the perch. Anolis bimaculatus hits the rocky ground and begins to sprint toward the nearest bush. At this moment a large wave crashes over him and carries him off the low-lying island. Anolis bimaculatus scrambles onto a floating log, but is immediately caught in the outgoing rip and is carried far into open water within moments. Anolis equestris watches A. bimaculatus ride the waves out of his life and weathers the remainder of the storm safely on his elevated perch, alone. ***Anolis equestris wins***
Which brings us to our dramatic conclusion: Anolis frenatus, representing the mainland, versus Anolis equestris, representing the islands!
The third round of our (fictional) anole tournament is now complete. We started with 32, and now there are 4.
Here’s what you missed in the Elite 8 round: Anolis aquaticus proves he’s in it to win it when he battles Anolis limifrons; Anolis proboscis wonders what his proboscis is good for as he battles the giant Anolis frenatus; Anolis equestris shows Anolis porcus who the king of the jungle is; and the big-cat-lizards Anolis marmoratus (AKA Leopard Anole) and Anolis bimaculatus (AKA Panther Anole) face off.
Our (fictional) tournament of anoles is well underway as we complete the Sweet 16 round! The competition was tough and a few favorites fell from glory (sorry!). We’ve been repeatedly asked how the competitions are being decided. We are not rolling dice or using random outcome generators. For each match, we (K. Winchell, M. Muñoz, and P. Muralidhar) are reading up on the biology of the species involved and we debate what would happen if they were to meet each other and try to highlight some facts about each. In cases where we might be biased we step aside and let the others decide. If it were up to me of course, Anolis cristatellus would be in the final! (Really, they’re scrappy and mean little guys!)
And now the results!
Anolis cristatellus on a smooth, vertical substrate in Puerto Rico (photo by K. Winchell)
In urban areas, the number of natural substrates (e.g. trees) is reduced. In their place are novel manmade substrates (e.g. walls, metal gates). These surfaces undoubtedly have different properties relevant to anole locomotion: they are smoother, harder, and (in the case of walls) much broader and flatter compared to natural surfaces in a forest. In urban areas lizards still use these substrates at high frequency, but do they do so effectively? Kolbe and co-authors began to dive into this complex topic in their recent publication, “City slickers: poor performance does not deter Anolis lizards from using artificial substrates in human-modified habitats” (Kolbe et al. 2015).
The relationship between habitat use, morphology, and performance for anoles has been extensively studied in natural environments (reviewed in Losos 2009). Urban environments add new dimensions to this area of research. Resource distribution and abundances differ drastically compared to natural areas. For example, the distribution of available perches and what they are made of in urban habitats is very different from a forest. Moreover, the properties of these resources differ drastically as well: urban substrates are smoother, broader, and have different thermal properties, to start. Understanding these differences in habitat use and how they influence performance and, ultimately, adaptive responses in anoles is the topic of ongoing collaborative research that I (K. Winchell) and the Kolbe lab have been conducting.
If you’ve been following along with our “Anole March Madness” tournament and you didn’t catch the updates on Twitter, you’re in luck. Here are the recaps of the second half of the Round of 32.
And of course, we remind our readers that these battles are fictional and are meant to highlight ecology, distribution, and traits of some well-known and not-so-well-known anoles. All battles are written by myself (K. Winchell), Martha Muñoz, and Pavitra Muralidhar using “complex algorithms” to determine the ultimate champion.
The next round (Sweet 16) kicks off on Twitter March 28 at 8pm. Follow along live and tag your comments #AnoleMM2016! Also next week we will be highlighting the “state of the knowledge” about the anoles featured in our tournament. Any guesses (no cheating) which species has the most citations and which has the least? We’ll give you the run-down of what we know and don’t know about who.
And here’s the recap of the second half of the Round of 32!
For those of you following the first ever Anole March Madness, the first eight matches were held on Monday and Tuesday evening live on twitter, taking us halfway through the round of 32!
Just a reminder, no animals were harmed in the making of this production and all matches are entirely fictional. We have done our best to make the battles realistic given what is known about the natural history of the species and their habitat. Of course, we have taken minor creative liberties with some details. We also want to point out that while these matchups are for entertainment, we hope you also learn a little about some of the more obscure anoles. It is also an opportunity for us to point out how little is known about some of these species. Please let us know in the comments if you believe some detail to be inaccurate or you have something to add!
And now, the recaps!
Day 1 was exciting with Anolis insignis versus Anolis nebulosus, Anolis onca versus Anolis macrolepis, Anolis cristatellus versus Anolis pulchellus, and Anolis lividus versus Anolis richardii.
Anolis insignis versus Anolis nebulosus
High in the treetops of Mexico perches the diminutive clouded anole, Anolis nebulosus. A loud rustling of leaves announces the arrival of the massive crown-giant, Anolis insignis. The lizard, famished from his travel from Panama, spots the tiny A. nebulosus and decides he looks like a tasty snack. He slowly ambles across the canopy towards his meal, pauses, then lunges with mouth agape. The agile A. nebulosus leaps to the side, narrowly avoiding the cavernous mouth. Anolis nebulosus watches with curiosity as the striped giant slowly turns toward him, preparing his next attack. Again, A. insignis lunges towards his hopeful meal and again A. nebulosus narrowly avoids being consumed. Wary of the newcomer, Anolis nebulosus retreats below the canopy, to the lower branches where he spots an attractive female. Foolishly thinking he has escaped the unknown giant in the tree top, he begins to dewlap at the prospective mate. Meanwhile, the hungry Anolis insignis stealthily descends to the lower branches and lunges again at the distracted A. nebulosus. This time he doesn’t miss and consumes the tiny anole in a single gulp. ***Anolis insignis wins***
Anolis onca versus Anolis macrolepis
At the terminal end of the babbling brook, the habitats of two Colombian anoles, Anolis onca and Anolis macrolepus, converge. Following an escaped beetle downstream, Anolis macrolepis leaps from rock to rock over the rushing water. On his last leap he finds himself no longer on a rock, but on soft sand. Here his troubles are much bigger than an escaped meal as he meets the larger Anolis onca, basking in the sun on the sand. Charged full of energy, A. onca confronts the intruder. Anolis macrolepis attempts to turn to flee, but his feet are poorly adapted to run on this strange, hot substrate. He moves his legs frantically attempting to run, but only digging himself into a hole. Now half buried in the sand, A. macrolepis realizes he is in a precarious situation. Anolis onca approaches his competitor and flings sand in his face. Temporarily blinded, A. macrolepis panics. He begins to flail out of desperation as A. onca firmly grabs him and flings him back into the stream where he is carried out to sea. Anolis onca resumes basking in his sandy home. ***Anolis onca wins***
Anolis pulchellus versus Anolis cristatellus
In a patch of secondary tropical forest on the north coast of Puerto Rico, Anolis cristatellus and Anolis pulchellus both enjoy the afternoon, each happily unaware of the other as they occupy distinct structural habitats. The afternoon sunshine begins to wane and A. pulchellus climbs up the side of a tree trunk towards one of the remaining patches of sunlight. Anolis cristatellus suddenly darts around from the other side of the trunk to confront the trespasser. Anolis cristatellus raises his dorsal crest and stands strong and proud, seeming to grow by almost 2 cm. Anolis pulchellus, with his eye on the prized sunlight, accepts the challenge. He too raises a dorsal crest, dark spots suddenly appear behind his eyes, he sticks his tongue out, and he flashes his large crimson dewlap with confidence. Unfazed, A. cristatellus in turn flashes his bright orange and yellow dewlap and does several pushups as a last warning. Anolis pulchellus does not heed the warning and darts toward A. cristatellus, delivering a sharp blow to his midsection with his pointy snout. Despite the sting of the assault, A. cristatellus manages to bite onto the tail of A. pulchellus. Unable to autotomize his tail, A. pulchellus is swiftly flung from the tree trunk, flying several meters back to his rightful place among the reeds of grass where he decides it prudent to remain, defeated. ***Anolis cristatellus wins***
Anolis lividus versus Anolis richardii
On the tiny isle of Montserrat, Anolis richardii has arrived on a piece of driftwood to make its stand against Anolis lividus. There are perches and food for the taking, and only a puny mid-sized anole to defend them. Anolis lividus has spotted this unwelcome intruder and rushes up the tree to evict him. Unaware of their shared evolutionary history, they are briefly confused when they flash their orange dewlaps and, for a moment, wonder whether they’re the same species. Anolis richardii, massive to begin with, cuts an even more impressive figure with an erected nuchal crest, but A. lividus does not back down. However, the Montserratian anole is no match for the massive A. richardii. Its lunges and parries are met with vicious attacks. Anolis richardii delivers a strong bite to A. lividus’ flank. At that moment, the Souffriere Hills volcanic dome collapses, launching a column of ash and pyroclastic material into the air. Anolis lividus is unfazed by the volcano, but A. richardii is terrified by this ominous neighbor and decides to find fertile ground elsewhere. It abandons the perch and the island, leaving A. lividus in disbelief at his luck and the unexpected victor of this encounter. ***Anolis lividus wins***
Day 2 was equally exciting with Anolis limifrons versus Anolis humilis, Anolis frenatus versus Anolis transversalis, Anolis placidus versus Anolis distichus, and Anolis trinitatis versus Anolis pogus.
Anolis limifrons versus Anolis humilis
It is early morning in the Costa Rican rainforest. Anolis limifrons has been awake for an hour already, foraging in the leaf litter near the towering buttress roots of a tree. As it plucks a roach into its mouth it freezes. There, not one meter away, is the red and yellow flash of the dewlap of Anolis humilis, a much bigger species. Anolis humilis charges, vying for the insect-rich territory. Anolis limifrons retreats to a crevice in the roots, but then charges back out and and lunges at A. humilis. Anolis humilis is stockier, but slower and less tenacious. Anolis limifrons won’t give up, biting at A. humilis often, displaying, and refusing to back down. Anolis humilis gives one last lunge, but misses, as A. limifrons deftly dodges and swings back around to nip his opponent’s throat. Harangued and weary, A. humilis retreats, leaving A. limifrons to go back to his cockroach feast. ***Anolis limifrons wins***
Anolis frenatus versus Anolis transversalis
It is early morning and Anolis transversalis, wide-eyed and nervous, glances around. There are so many things that can eat him here in Amazonia. From above comes an Anolis frenatus, crashing down from the canopy. The crown-giant, startled by his misstep and fall, barely grasps the tree branch as he falls. Swinging upright, he notices he is not alone. Anolis transversalis, petrified that his worst nightmares have come true, turns to run. Anolis frenatus, still confused from the fall, watches as A. transversalis flees frantically, jumping from the branch to an adjacent tree, then to another and another and another until he is out of sight. Uncertain about what just occurred, A. frenatus unknowingly claims victory. ***Anolis frenatus wins***
Anolis distichus versus Anolis placidus
It’s late evening now on Hispaniola and the placid anole, Anolis placidus, peacefully perches high in the treetop hugging the twig he plans to sleep on tightly with his tiny legs. Anolis distichus, thinking it best to sleep higher up tonight, ascends to what looks like the best sleeping site only to find it occupied by the little twig anole. Anolis distichus, feeling entitled to this perch, strolls up to A. placidus and begins to display his dewlap. Anolis placidus, ever the pacifist, looks at A. distichus with a sidelong glance and decides there is room enough for them both. Anolis placidus sidles to the end of the perch, pressing his body close to the twig. Anolis distichus, upset that his opponent does not accept his challenge, advances and begins to push-up violently, thinking he may shake the twig anole off of the branch. Anolis placidus edges closer to the end, his tiny legs giving him an unshakeable grip. Clearly his friend needs more room to sleep, that’s fine with him, they can share. Anolis distichus, frustrated with his opponent and now struggling to keep balanced on the moving twig decides intimidation is not enough and leaps towards A. placidus, who skillfully moves to the underside of the twig with only two steps. Anolis distichus lands on the edge of the twig but is unable to grip the tiny perch with his longer limbs. As he flails attempting to grab onto the twig and his opponent, A. placidus inches back away from the edge to the top side to avoid contact. Anolis distichus manages to catch the tip of the branch with a single toe, momentarily hanging on before falling to the branch just below. As his toe releases it turns the twig into a catapult and even the powerful grip of A. placidus isn’t strong enough to hold on. Anolis distichus looks up as A. placidus is flung far out of sight. Anolis distichus ascends to his chosen sleep site, pleased with his skillful removal of the competitor. ***Anolis distichus wins***
Anolis pogus versus Anolis trinitatis
Aboard floating vegetation following a hurricane that tore through the Northern Lesser Antilles, Anolis pogus patiently awaits making landfall on St. Vincent island. As the flotsam approaches the shore, the weary castaway spots movement in the trees. Flashes of bright green and blue announce the presence of the resident trunk-crown anole, Anolis trinitatis. Anolis pogus disembarks on the rocky shore and wearily climbs up the nearest trunk, basking in the sunlight and surveying his new home. But there will be no rest for the tired traveller, immediately an A. trinitatis descends and sizes up this newcomer. Anolis trinitatis, a good 2cm larger in body size, decides he does not like new, strange looking anole. He angrily advances and grabs A. pogus by the nape of the neck and flings him. “Welcome to St. Vincent” he thinks, hoping A. pogus will choose somewhere else to call home. Suddenly energized, A. pogus climbs up the adjacent tree and begins to dewlap at his assailant, undeterred by his aggression and larger size. Anolis trinitatis leaps to the tree, accepting the challenge. They trade blows repeatedly – A. trinitatis bites at the face of A. pogus; A. pogus leaps and bites off a sizable portion of A. trinitatis’ tail – both sustain substantial injuries in the battle. Now weary himself and impressed with his opponent’s stamina and scrappiness, Anolis trinitatis attempts one last time to disarm his opponent with intimidation, he dewlaps and pushups but his vigor is gone. The bloodied A. pogus returns the threat by advancing and delivering a calculated blow to the throat, tearing the dewlap of A. trinitatis. Anolis trinitatis has had enough and limps away, defeated, back to the canopy to inform the other native lizards that there is a new lizard on the block and that he’s not one to be messed with. ***Anolis pogus wins***
Tune in tomorrow and friday on Twitter for the next matches, and of course we will bring you the recaps here afterwards. Bracket busted already? Did we miss an important detail? Let us know in the comments!
Get ready! Anole March Madness begins with the first battles this evening (Monday 3/21/2016) . We will be live broadcasting the bouts on twitter with the handle @AnolisMM2016 starting at 8pm EDT (Boston time). Tonights matches include: A. insignis v. A. nebulous, A. onca v. A. macrolepus, A. cristatellus v. A. pulchellus, and A. lividus v. A. richardii.
Get your brackets filled out, share them on twitter with #AnoleMM2016, and follow along with the live tweets. We will also be posting the recaps of the battles tomorrow morning here on Anole Annals!
Let the games begin!
Are you ready for March Madness?! No, not the basketball tournament… this year we are staging the very first Anole March Madness! We should acknowledge that we came up with this idea after following the popular Mammal March Madness for the past couple of years and feeling a little left out.
What is Anole March Madness? Just like the basketball tournament, we start with 32 of the best teams from 4 regions. The groups doing battle will be Mainland (Central America and South America) versus Island (Greater Antilles and Lesser Antilles). The rules are simple, if a lizard wins a bout it advances, if it loses it is out of the tournament.
Anole-March-Madness: download this pdf, fill in the text boxes, play along!
Don’t worry, no animals will be harmed in any way during this tournament. In fact, the matches are entirely fictional and come from the imaginations of AA contributors Kristin Winchell, Martha Muñoz, and Pavitra Muralidhar. We will pick the winners of each bout based on what is known about the morphology, ecology, and behavior of each species and will give a full recap of the (fictional) battles every few days.
Do your best to predict the winner of each match, and ultimately the champion! Fill in the empty text boxes in the PDF of the bracket above and pick your winners for each match. Take a photo of your completed bracket and tweet it with #AnoleMM2016 so we can all see your picks! You earn 1 point for each correct match in the first round (round of 32), 2 points in the second round (Sweet 16), 4 points in the third round (Elite 8), 8 points for the fourth round (Final 4), and 16 points if you guess the champion. Keep score and let us know how you’re doing! First matches will be posted in a few days. The Championship match will be on April 4th.
Now, meet the teams!
An actual perch used by Anolis cristatellus – neither smooth nor flat! (photo by K. Winchell)
In the real world, lizards cling to everything from smooth, flat concrete walls to rough, rounded tree trunks. So why is it that most studies on cling force in anoles focus on clinging to smooth flat substrates? Does cling force differ if the substrate is rounded or rough? Jason Kolbe sought to answer this question in his recent publication, “Effects of Hind-Limb Length and Perch Diameter on Clinging Performance in Anolis Lizards from the British Virgin Islands” (Kolbe 2015).
We know that morphology impacts performance in anoles and that performance varies with environment. For example, sprint speed is correlated with limb length, but this relationship depends on the diameter of the substrate (e.g. Losos and Sinervo, 1989). We also know a little about clinging performance in anoles. Greater cling force is correlated with larger toepads and more lamellae on smooth flat surfaces (Irschick et al., 1996; Zani 2000; Elstrott and Irschick, 2004), but adhesion on rougher surfaces may be influenced by claw and toe morphology (Zani 2000).
There appears to be an unexplored interaction between substrate properties and clinging ability that involves more than just toepad characteristics. Specifically, Kolbe points out that claws can increase clinging ability by digging into the perch or simply by increasing friction on the surface. Limbs can also increase friction via the application of compression forces to the substrate. In other words, cling force can be increased, particularly on rough surfaces, by using muscular force to grasp rather than relying on van der Waals forces from the toepads, which are more effective on smooth flat surfaces.
Anole species used in this study: Anolis cristatellus (left) and Anolis stratulus (right). Photos by K. Winchell.
Kolbe investigated this further by looking at the interaction between limb length and clinging ability on perches of different diameters with Anolis cristatellus and Anolis stratulus from the British Virgin Islands. Specifically, he hypothesized that cling force should increase as the ability of a lizard to obtain a firm grasp on a substrate improves (i.e. when it can wrap its limbs around the substrate). This ability to form a secure grasp is dependent on both the diameter of the perch and on lizard limb length.
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