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W
e just wrapped up our Anole March Madness tournament in which we brought to you many fictional battles of mainland and island anoles. We researched each species included in the tournament and highlighted some little-known facts about the participants. We told you about dewlap colors, ecomorphs, habitat use, invasion, body patterns and colors, and much more. But finding this information for each species was more challenging than we originally expected. The state of anole knowledge is certainly imbalanced.
Looking at the number of Google Scholar citations for each species shows just how imbalanced our knowledge of different species of anoles is. Of the species we included, only three of the mainland species had more than 200 citations and none had more than 600. In contrast, of the island species, six had more than 200 citations and Anolis cristatellus had a whopping 1230 citations! Within regions there is quite an imbalance, too — check out how many more studies there are on Central American species compared to South American species (orange and red) and on Greater Antillean species compared to Lesser Antillean species (light and dark blue). This is perhaps unsurprising to many. We all know that anole research tends to be biased towards Caribbean species, but it was surprising to me how large this bias is!
Google Scholar hits for mainland species in Anole MM 2016
Google Scholar hits for island species in Anole MM 2016
This gives a sense of how much information is available for the species we highlighted in our tournament. For a few species, we were not even able to find photos (at least with appropriate usage rights) and for many the only information we could find was native range, color, and body size. For example, Anolis placidus had only five Google Scholar citations to go by, few images (other than specimens), and really no ecological information other than its ecomorph. But it must be named the “Placid Anole” for a reason, right? So we ran with that. Even with the paucity of information for some species, we were able to piece together some interesting stories highlighting some of the unique adaptations, ecology, and morphology. Here are some of my favorites with links for more information.
Anolis aquaticus (photo by John Sullivan)
Anolis macrolepis (photo by fernandocastro)
Anolis aquaticus and Anolis macrolepis were our two representative aquatic anoles. These species are adapted to streamside living, using the boulders as foraging grounds and for quick escape from predators–swimming, running across water, and even remaining submerged to get away from a threat! We used these behaviors in our stories–allowing both species to have sure footing in their stream side home and escape into the water when necessary.
The padless foot of Anolis onca. Photo by J. Losos.
Another unique anole we featured was Anolis onca, the only anole to have lost its toepads entirely. Anolis onca is adapted to a beachy life, making its home in sandy substrates and perching on bushes. We played this as a strength in round 1. It’s not clear what the functional reason for the loss of lamellae is–as has been suggested on Anole Annals, perhaps the sand renders lamellae or setae useless. The reasons for this morphological outlier are yet to be explored, but we figured A. onca must have an advantage in the sand over other anoles!
Anolis proboscis (photo by Luke Mahler)
Of course we had to feature the Anole Annals darling, Anolis proboscis. Here at Anole Annals we’ve covered everything from babies hatching with tiny horns to discussion of their odd looks to the “rediscovery” of this odd Pinnochio anole. What exactly that horn is for seems to be up for some debate still, but it seems that it is likely involved in sexual selection: male-male competition or female choice. And the part in the battle where the anole wiggles his snout? Yep, they do that!
Anolis gorgonae (photo by Luke Mahler)
A crowd favorite from our tournament, but alas not the champion, was the only completely blue anole: Anolis gorgonae. This anole didn’t make it too far in our tournament because his bright blue body made him extremely visible on the forest floor. The blue may help camouflage them way up in the canopy, but it seems that down low they are visible against the green leaves from a distance. We wanted to root for him too, but how does something that visible not get eaten to extinction?
Anolis insignis (photo by Shawn Mallan)
Anolis frenatus (photo by Tobias Eisenberg)
Anolis equestris, high in the canopy (photo by Corn Farmer – Flickr)
Anolis garmani (photo Wikipedia)
Anolis richardii (photo Wikipedia)
Anolis bimaculatus (Photo by Statia Wildlife, Flickr)
And finally, we made sure to feature a few of the anole kings: the crown giants. We know our readers here at Anole Annals love their giant anoles. These monsters of the canopy seem like they could beat out just about anything (or eat them!), and so it was in our tournament. In the end it was too tough to decide between the beloved Anolis equestris and the lesser-known giant of the mainland, Anolis frenatus. And so we enlisted the help of the mainland-island competition vs predation debate (reviewed in Losos 2009 – p.159) and had a monkey swoop in to help us decide (yes, this happens!). We wondered, could the predator-naive A. equestris establish itself in a land full of predators like the forests of South America? Despite being successful at invading Miami and some other Caribbean islands, it doesn’t seem to have established itself in South America… yet.
On behalf of all of the contributors to Anole March Madness 2016 we thank you for playing along with us. We hope to bring you another tournament next year, so let us know if you want to help make that happen!
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!
Anolis frenatus (photo by Tobias Eisenberg)
Anolis equestris (photo Wikipedia)
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:
Anolis aquaticus (photo by John Sullivan)
Anolis frenatus (photo by Tobias Eisenberg)
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***
Anolis equestris (photo Wikipedia)
Anolis bimaculatus (Photo by Statia Wildlife, Flickr)
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.
I am very interested in IDing the following anoles I saw on the island of Culebra, which is 17 miles east of Puerto Rico. I thought I would be able to find a guide to anoles of this area online, but IDing them turned out to be harder than I thought. Thank you for your time!
1) First one was found in the parking lot of Flamenco.
2) Also found near Flamenco.
3) Found in town. Bright yellow dewlap.
4) Found in town.
5) Found in town.
6) Found in town. Dewlap was dark orange with yellow near the throat.
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!
A few days ago, I had a very interesting discussion in twitter with some colleagues (see here [1] and here [2]) about whether there is a decline and crisis of new taxonomists in amphibians and reptiles. I wonder if this situation might be the case fir Anolis lizards, or whether Anolis has its own tendency in species description. So, I decided to take the bull by the horns and I went to the Uetz Reptilia Database to see the numbers.
First, it seems that there is not an increase in species description for anoles across time. In early years, it was a bunch of new descriptions as also recently (Fig 1).
However, if you check how many anole taxonomists have described species, an interesting pattern emerges. Very few people have described almost all of anole diversity (Table 1). Only 15 anologists have described all currently known species (400 according to the Uetz database), and more interesting is the fact that only five of them are still alive! (Gunther Köhler, Orlando Garrido, James McCraine, Steven Poe and Larry Wilson; Table 1).
Table 1. Ranking of anole taxonomists by number of species described.
Then, to establish if new authors (new anole taxonomists) are emerging in recent times, I made an accumulation curve of authors across years of description (Fig. 2). I considered only the senior author (Fig. 2, top) and all authors involved in the description (Fig. 2, bottom). From these figures, it is possible to see, apparently, that very few new authors are emerging as anole taxonomist specialists.
Accumulation of authors by year. Only senior author (top), all authors involved in the description (bottom)
This suggests, at least to me, that something is happening in anole taxonomy. As we discussed in twitter, this scarcity in taxonomists in recent times could be due to several factors: lack of interest in describing new forms, lack of funding resources, lack of access to comparative material (i.e., herpetological collections), lack of writing skills to generate scientific papers, or lack of adequate taxonomic training. In any case, I feel that anole taxonomy would be in crisis if new people, mostly from Latin America, are not interested in describing and revising new species in each country. But, why are very few people seem to be interested in describing new valid species nowadays? Is it perhaps that taxonomy is a discipline with little interest in academia? Does doing so serve to get a job? Is it not worth publishing papers in taxonomy or even describing new forms?
Although today there are a bunch of journals oriented to taxonomy and many online resources to access to primary literature (Sci-Hub [3], BHL[4], etc.), it seems that very few people are interested to tackle these problems in our iconic lizards.
I would like to see your opinions on this
References
[1] https://twitter.com/CrawfordAJ/status/713398834005561344
[2] https://twitter.com/CrawfordAJ/status/713459231983280130
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!
From Skip Lazell’s 1972 monograph
Like many other biodiversity journals, the contents of the Bulletin of the Museum of Comparative Zoology are available on the Biodiversity Heritage Library. However, the BHL can be somewhat cumbersome to deal with, if not downright counterintuitive. Just today, I downloaded Lazell’s spectacular, classic monograph on the Lesser Antillean anoles, full of detailed descriptions, lovely illustrations (as above) and incisive commentary. Every digital library should have a copy, and so here’s your chance to get one easily, by clicking on this link. But note: the link apparently is only good for 30 days (and someone had to try it twice to get it to work, so be persistent). Also note: it’s a big file, so be patient as it downloads.
Enjoy! And after you look it over, write a nice comment for Skip, AA‘s all-time leading commenter, to read.