Author: Adam Algar Page 1 of 2

Associate Professor at the University of Nottingham (UK). My research focuses on niche limits and dynamics across scales, from individual organisms to the globe. Mostly lizards.

Some would say there is a glorious history of anole-inspired poetry; others would say there is a history. But everyone can agree that it has been brief. Regardless, the best—and worst—of this work has been featured on Anole Annals (and nowhere else*). Last week, with multiple looming grant and paper deadlines, plus a lecture (or two or three) to write, I didn’t anticipate adding to this history (after reading the below, you’ll be glad that it is, now, history). However, I did open Twitter which is a pretty good sign that I was running out of useful ways to procrastinate. Near the top of my feed was this tweet/reply combo from Ken Feeley and James Stroud:

Without question, these are both great titles, but they brought another to my mind:

“Anolis, how do I love thee?”

Having begun, and with procrastination as my muse, I couldn’t stop there and leave the thing unfinished. But, before you read further, if you’re not familiar with the famous poem whose title I have modified (i.e. ruined), you can read it here. It may help make sense of what follows (or maybe not). Regardless, to the memory of Elizabeth Barrett Browning and all appreciators of beautiful (or any other kind of) literature, I am deeply sorry.

Anolis, how do I love thee? Let me count the ways.
I love thee to the depth and breadth and height
My lasso can reach, when clinging out of sight
For the ends of twigs and trunk and grass.
I love thee to the level of every day’s
Sampling size, by sun and head-light.
I love thee freely, as we strive for statistics right.
I love thee purely, as they remain robust.
I love thee with the passion put to use
In my old hypothesis, and with my co-author’s faith.
I love thee with a love I seemed to lose
With my lost data. I love thee with the breath,
Smiles, tears, of all my life; and, if the AE choose,
I shall but love thee better after the reviews.

So there it is. You may tolerate it or hate it. No on agrees on literature (very loosely defined) anyway. However, if there’s one thing we can agree on, it is that the whole thing is James Stroud’s fault (just kidding, James, I take full responsibility).

I posted the ‘poem’ on Twitter and so it is only fitting that Twitter should have the last word. Thus, I’ll leave you with this reply to my tweet from one of my past PhD students, which I think sums up the whole experience rather nicely:

*This isn’t actually true, but hey – artistic license. It turns out anole poetry has featured elsewhere and the history hasn’t been that brief either: https://www.anoleannals.org/2012/07/11/anole-poetry/

Forget Brexit, It’s BrAnolis: An Anole Makes the Cover of the British Ecological Society’s Bulletin Magazine, But What Species Is It?

By Adam Algar

On April 12, 2018

In All Posts, Anole Photographs

A Costa Rican anole graces the cover of the March 2018 issue of the British Ecological Society’s magazine ‘The Bulletin’. But what species is it? Photo by Roberto García Roa.

When you think of hotbeds of Anolis research, the United Kingdom probably isn’t the place that immediately springs to mind. And unsurprisingly – there are no dewlaps decorating the tree trunks of Wytham Woods. Of course, there is a strong tradition of anole research on this side of the pond including Roger Thorpe and Anita Malhotra at Bangor University and Katharina Wollenberg-Valero at the University of Hull (there’s undoubtedly more, past and present, of which I’m not aware; sorry if I’ve left you out!). Still, we’re unlikely to be hosting a spin-off Anolis Symposium (Anolis SympX?) any time soon and anoles certainly don’t dominate British Ecological Society (BES) meetings like they do SICB or Evolution. So, needless to say, when the March issue of the BES’s members’ publication The Bulletin thunked through my mail slot a few weeks ago, I was surprised to see one of this blog’s titular reptiles staring back at me. Though maybe I shouldn’t have been surprised. After all, we know that an anole loves a good cover shot. A quick delve into the magazine revealed that the photo was taken by Roberto García Roa from the University of Valencia and that Roberto won the Up Close and Personal category in the BES’s photography competition for his ‘mid-shed’ shot.

Roberto García Roa’s award winning anole photo.

My question for the Anole Annals readership is this: can anyone identify the species? The photo was taken in Costa Rica, but there’s no more information than that. Any thoughts?

Editor’s Note July 3, 2018: Robert Garcia Roa has provided this photo, which reveals that the species is <i>Anolis cristatellus</i>, an invasive species in Costa Rica.

The Contronymic Effect of Behaviour on Evolution: Does Behaviour Promote or Retard Evolutionary Change?

By Adam Algar

On November 22, 2017

In All Posts

Anolis shrevei on a rock. Photo courtesy of Katharina Wollenberg Valero.

Back when I was an undergrad, we were given the impression that animal behaviour and ecology (and evolution for that matter) were distinct disciplines. ‘Behaviour’ had its own classes, professors, and students, mostly separate from the ecologists. The disciplines also have their own, aptly named, journals: Animal Behaviour vs. Ecology. Of course, even then we knew that this division wasn’t a hard boundary and journals like The American Naturalist explicitly included ecology, behaviour and evolution, but we still didn’t think of these disciplines as inseparable. The impression seems to have stuck because fast forward 15 years and I was recently surprised when a colleague, upon hearing about our work on microhabitat use in agamids said, “I didn’t know you did behaviour.” Well, until that moment, I didn’t know I did either! I simply hadn’t thought about what we were doing in that way. Upon reflection, it should have been obvious, just like it should have been obvious that the behaviour/ecology distinction was a false one. I have no idea how widespread my once-perceived separation of ecology and behaviour is, but the fact that there’s a need for a journal, Behavioural Ecology, that specifically merges them suggests they’re still not perfectly integrated.

So why natter on about behaviour, ecology and evolution? Because a recent paper by Martha Muñoz and Jonathan Losos, published in The American Naturalist, is a fine example of why these shouldn’t be separated. Muñoz and Losos set up a dichotomy of hypotheses about how behaviour influences evolution: on the one hand, exploratory behaviour can expose species to novel selection pressures, stimulating evolution, but on the other, behavioural fidelity could shield species from those same selection pressures, ‘forestalling’ evolution. So, which is it? Well—spoiler alert—it’s both. So long nice dichotomy. To reach their findings, the authors looked at thermoregulatory behaviour and how it affects adaptation to high elevation habitats in the Anolis cybotes species group (specifically cybotes, armouri and shrevei). They found that, despite much cooler temperatures at higher elevations, high and low elevation species had selected temperatures in the lab and maintained similar body temperatures in the field, via increased thermoregulation at high elevations. Thus, despite the cooler temperatures, anoles hadn’t evolved to prefer colder temperatures on mountaintops. So behaviour halts evolution, right? Yes but no. To thermoregulate so extensively, anoles had to seek out warmer microhabitats, specifically boulders. And we know what happens when anoles change their perch type: evolution! Muñoz and Losos found that shrevei and armouri had flatter skulls, consistent with life on the rocks, as well as shorter hind limbs (but no differences in toe length or lamellae number). The evolutionary basis of the morphological change in head and femur traits was confirmed by a common garden. Nifty.

Effects of behavioral thermoregulation on evolution of high elevation anoles. On the left, thermal environment, body temperature and lab-selected-temperature of low and high elevation anoles. On the right, morphology of high and low elevation anoles in the field and in a common garden. Modified from Figs 1 and 2 in Muñoz and Losos (2018).

The overall message of the paper is clear: the same behaviour inhibited evolution along one niche axis and promoted it along another. Muñoz and Losos argue that the lack of evolutionary change in thermal traits arises from the Bogert effect, where behaviour limits exposure to novel selection pressures. However, there is a chance that the lack of evolution could be due to other constraints, like a lack of genetic variation. Testing this would require an experiment with a control group of lizards that couldn’t behaviourally avoid thermal selection pressures. A previous paper by Muñoz, Losos and others, provided just such a natural experiment. In that study, Muñoz et al. found that lower CTmin has evolved in high elevation cybotoids, relative to low elevation ones. Why? Because at night, when its coldest, anoles are unable to behaviourally thermoregulate to avoid the cold –voilà, a control where the Bogert effect was negated. And once behaviour was removed from the equation? Evolution! This finding adds even more weight to the role of behaviour in inhibiting the evolution of thermal traits in this system. Cool stuff (I make no apologies for that pun).

There’s lots more in the Muñoz and Losos paper than I touched on here so give it a thorough read. It goes a long way to destroying any divisions that might still exist between behaviour, ecology and evolution and it makes a strong case for why we need to consider multiple niche dimensions when we talk about niche evolution and conservatism. Plus, it gave me an excuse to use the word ‘contronymic.’

Muñoz, M.M. & Losos, J.B. 2018. Thermoregulatory behavior simultaneously promotes and forestalls evolution in a tropical lizard. The American Naturalist. DOI:10.1086/694779.

Video of Running Trunk-Ground Anole Needed

By Adam Algar

On November 15, 2014

In Anole Videos

I’m looking for a bit of help and where else to turn than the dedicated readers of Anole Annals? Does anyone have a short video clip (ca. 10 seconds) of a trunk-ground anole running on either the ground or a trunk that they’d be willing to share? I’d like to use it for a couple of upcoming talks, and for teaching. Proper credit would, of course, be given. Plus I’ll buy you a beer if you ever happen to be in Nottingham. I’ve got a few short clips of sagrei but unfortunately the frame rate went screwy when I tried to convert them, hence the appeal. The point is to contrast a trunk-ground’s movement with this clip of carolinensis (shot by Leslie Bode on the Anhinga Trail, Everglades, FL):

If you have something suitable that you’re willing to share, please either leave a comment, or you can email or tweet me (adam.algar[at]nottingham.ac.uk, @acalgar).

Thanks!

Globalization and the 50-Year-Old Predicted Reorganization of Anole Biogeography

By Adam Algar

On September 27, 2014

In New Research

“I caught an anole lizard and tossed it ten feet or so out into the water. To my dismay, it popped to the surface, swam expertly back to the shelter of the trees, and climbed up a mangrove trunk. Well, I continued, suppose a full hurricane blew an anole so far away on open water it couldn’t get back. Our little experiment shows that it could swim to the nearest islet if it were not too far away.”

So wrote E.O. Wilson (1995 p. 271, Warner Books, NY) in his autobiography, Naturalist, reflecting on his island defaunation work with Daniel Simberloff. From his ‘little experiment’ (I can hear animal care committees cringing), Wilson postulated that anoles could, if they had to, disperse from island to island across open water. Whether anoles can cross water, however, isn’t that important. Rather, what’s important is that they rarely do. Anoles’ status as a symbol of island biogeography and adaptive radiation is largely due to the fact that isolation and the resulting low gene flow among islands set the stage for in situ speciation and adaptive radiation. In fact, much of what we (and island biogeography in general) owe to anoles, we owe because they don’t swim so well. And they don’t colonize new islands very often.

Or rather, they didn’t.

A new paper in Nature by Matt Helmus, with AA stalwarts Luke Mahler and Jonathan Losos, shows how human-mediated dispersal of anoles among Caribbean islands is reorganizing anole biogeography in a very predictable way. I suspect many who have worked on anole island biogeography, me included, have considered what to do about recent introductions and have often, like me, dropped them out of a dateset with the goal of trying to discern the ‘natural’ pattern. Helmus et al., however, saw the spate of recent anole introductions across the Caribbean as an opportunity, rather than a nuisance. Their great leap came from realizing that this reorganization of anole Caribbean biogeography should be predictable from the basic tenets of island biogeography theory.

Based on MacArthur and Wilson’s equilibrium theory, adaptive radiation theory and drawing on Losos and colleagues’ past work from 1993 and 2000, Helmus et al. predicted three patterns: 1) Species-impoverished islands (for their size) should have more exotics than more saturated islands, 2) The phylogenetic diversity of islands should increase due to exotic establishment, and 3) Human-mediated introductions should degrade richness–(geographic) isolation relationships. In short, they found evidence consistent with all of these patterns. Furthermore, they showed that economics that has replaced distance as the key determinant of island isolation. Needless to say, these are very exciting results that have supplied a key test, at biogeographic scales, of some classic theory*. It’s a must read.

This paper is also important because it shows how ‘blue skies’, curiosity-driven science can help us understand and, most importantly, predict how human activity will impact ecological systems. Did MacArthur and Wilson know, more than half a century ago, that their work would predict how increasing globalization and trade embargoes would affect modern biodiversity? I doubt it (cue someone pointing out in the Comments the exact line in the ETIB where they do predict this). However, regardless of whether they knew it at the time, this is exactly what their theory has done. As Helmus et al. state (p. 545): “Our results support the theory that it is the influence of geographic area and isolation on … speciation and colonization that fundamentally determine island biodiversity”. However, as they crucially find, what we now need to do is rethink how we define ‘isolation’. We can’t leave ourselves out of the equation any more. It’s economics, not geography, that matters now. Thus, not only does Helmus et al.’s paper test a long-standing theory, but it provides a clear example of the importance of fundamental scientific theory for understanding and predicting ecological dynamics in the ‘Anthropocene’.

In conclusion, the observation that humans are moving anoles — and other taxa — around faster than they could make their own way will come as a surprise to no one. But finding that the subsequent reorganization of life can be predicted by island biogeographic theory is fantastic (it should be pretty clear by this point that I like this paper. A lot). So if you haven’t read the paper, you should. I know it’s a terrible cliché to call a study ‘elegant’. So I won’t. I’ll call it damn elegant.

*I can’t help but mention that Helmus et al.’s findings were mostly based on good old-fashion OLS regression and ANOVA, and visualized using simple scatterplots – No fancy-shmancy statistical machismo here (phylogenetics aside). Just a clear set of predictions that could be parsimoniously tested. Chapeau.

Editor’s note #1: nice summaries of this paper have been written by Ed Yong’s Phenomena: Not Exactly Rocket Science blog and by Emily Singer in the new online Science magazine Quanta.

Editor’s note #2: The paper grabbed the cover of Nature.

This, in turn, joins a long list of recent science journal covers sporting an anole:

When the Going Gets Cold, Anoles Get Colder

By Adam Algar

On January 29, 2014

In All Posts

CTmax, Tb, and CTmin of cybotoid anoles & env. temperature. Modified from Fig 2 in Muñoz et al.

AA contributor Martha Muñoz’s work on altitudinal variation in the cybotoid anoles has already netted her the Raymond B. Huey award and of course, been featured on AA. A big chunk of this work, co-first authored with Maureen Stimola, has just been published by the Proceedings of the Royal Society B. If you haven’t read it yet, check it out.

I love this paper. However, in the spirit of full disclosure, I’m completely biased as I happen of be one of the co-authors. But I’m sure I’d love it anyway. Why? In part because it tests a clear hypothesis using multiple lines of evidence and eliminates confounding explanations – characteristics every paper should have. It also has cool (or should I say hot?) results. However, more than this, I think this paper demonstrates the power of combining good ole’ fashioned (yet cutting edge) field work with macroecological and macroevolutionary models, demonstrating how these different approaches can really complement each other.

What did Muñoz and company find? Briefly, they looked at hot and cold tolerance (CTmax and CTmin) of six species of cybotoid anoles on Hispaniola, in relation to elevation. They found far more variation in CTmin than CTmax across species (and populations). By bringing in a little macroecology, they showed that CTmax isn’t correlated with environmental temperature, but CTmin is, i.e. when the going gets cold, the anoles get colder – sort of. The catch is that while CTmin strongly tracks temperature, daytime body temperature does not. This is a neat result in and of itself and fits well with a big, recent, data-mining paper showing similar trends across hundreds of both ecto- and endothermic species. But while it doesn’t have the breadth of that paper, Muñoz et al. were able to go further. Firstly, bringing in a little macroevolutionary analysis, they showed that yes, CTmin has actually evolved significantly faster than CTmax. Neat, but at this point you should be asking yourself, “What about acclimatization?” and “Is this just plasticity?” Muñoz et al. asked the same thing and headed back to the field. A lot of work later and the answer was no. An acclimation experiment rejected this possibility.

At this stage, most macroecological and macroevolutionary analyses would have to stop at the identification of a clear, and intriguing pattern of fast past evolution of cold tolerance along an elevation gradient, but little CTmax evolution. The Discussion of such a paper would suggest potential hypotheses to explain the pattern and that would be that. But Muñoz et al. again went further and, by working in the field to measure perch use and operative temperatures, worked out why . The key result showed that lizards can behaviourally thermoregulate to escape the heat, thus reducing selection on heat tolerance, i.e. the Bogert effect. However, the nighttime cold cannot be escaped (actually, it can, by moving to England where it hasn’t dipped below -2 deg C this winter. Enjoy that polar vortex America!), leading to selection on cold tolerance.

Like I said, very cool results and a real testament to the power of using field experiments and macroevolutionary models to inform each other and go beyond what each approach could do in isolation. So please read it, challenge it, and build on it.

The Anole Effect In Taxonomic Research Bias

By Adam Algar

On November 16, 2013

In All Posts

Everyone (who reads this blog) knows that anoles are one of the most fascinating groups of organisms on the planet. We also all know that this makes them a remarkably popular lizard genus to study, a topic that Rich Glor has addressed on this blog previously. However, while dominant in some fields (and meetings, like Evolution or SICB) anoles scarcely make an appearance at others (ESA, IBS). This raises the question of whether anoles are really as popular as we think? How do they stack up to other sexy taxonomic groups like mammals, birds or bees?

A recent tweet and blog post by Chris Buddle from McGill gives the basis for an answer. He used a (quick and dirty) Web of Science search to find the number of publications per species within 15 haphazardly chosen Orders, merging these into higher taxonomic groups for visual comparison:

Originally from: www.scilogs.com/expiscor/biodiversity-bias-the-relationship-between-taxon-diversity-and-research-publications

Buddle showed that research publications are strongly biased toward mammals (no surprise there), then herps, then birds (I admit, a priori, I thought 2nd and 3rd would be reversed). However, the obvious question which jumped to my mind (and, I’m sure, to yours) was: how do our favourite dewlapping demons compare? A quick search on Web of Science with search string ‘Anole OR Anolis’ (yes I could have gone with anol*, but this picked up a few thousand extra hits for anolyte) gave approximately 12,686 hits. Using The Reptile Database‘s number of 393 anole species (cue argument now), I roughly mapped the anole results on to Buddle’s plot:

Modified from: www.scilogs.com/expiscor/biodiversity-bias-the-relationship-between-taxon-diversity-and-research-publications

Anoles win! Perhaps not surprisingly, anoles are remarkably well studied given their diversity, at least compared to other groups defined at the Order level (keep in mind that, because publication intensity is uneven within taxa, averaging across a more diverse group will tend lower the numbers). Nonetheless, anoles annihilate the carnivores, which Buddle highlights as the most studied order with a publication to species ratio of 7. However, let’s not forget that the vast majority of these studies are on sagrei or carolinensis, so there are still hundreds of under-studied anoles (and really, can a species be over-studied?)

Lastly, before we get too smug about the popularity of our exalted study genus, with its publication to species ratio (pub:spp) of 32.3, I did a quick look at the numbers for the genus Parus. This genus (even after lumping it in with Poecile, Cyanistes, Lophophanes, and Periparus) had a pub:spp = 327.1. And within the herps? Well, Sceloporus clocks in at 85.7. And Xenopus‘ pub:spp is 12451.3! Of course, Xenopus’ ratio is heavily influenced by biomedical research. Also, as I mentioned above, the uneven study of species within genera means averaging over a diverse genus like anoles will drag down the ratio, compared to less diverse genera like Parus, Sceloporus, or Xenopus. Even so, while anoles are clearly very popular, we undoubtedly still have a lot of work still to do.

WTF Anole Evolution?

By Adam Algar

On January 19, 2013

In All Posts

The website WTF Evolution has been popping up around the ecology and evolution blogosphere lately. It was recently featured by Jerry Coyne at Why Evolution is True and by Jeremy Fox at Dynamic Ecology. However, if you don’t read either of these blogs, or haven’t heard about the site, I suggest having a look – it’s pretty funny (especially ‘Fiddler crab’). Best of all, however, is that anoles are featured (and they’re the only organism to get a video)! The example is one featured on Anole Annals previously and is from a study by Casey Gilman in Duncan Irschick’s lab. The video shows that when anoles jump from springy perches, the recoiling perch can hit them in the tail, throwing them off balance. As the captionist at WTF Evolution notes, not a great thing for a lizard that spends a lot of time on branches: “Great work, Evolution“.

IBS Update: Sub-Fossil Anoles on Anguilla

By Adam Algar

On January 13, 2013

In New Research

Hello once again from the IBS in Miami. As I mentioned in my last post, anole presentations are a little thin on the ground here (though not around the conference centre where, with the help from some locals, we saw sagrei, carolinensis, equestris and distichus today).

I did have the opportunity yesterday to check out the the second of two anole presentations here – a new look at the sub-fossil lizard communities, with a focus on a question familiar to many readers of this blog: What’s the deal with A. pogus on Anguilla?

Has A. pogus ever been on Anguilla? Nope.

The study is being led by Melissa Kemp, from Liz Hadly’s lab at Stanford. Melissa has re-analyzed lizard dentaries from excavations on Anguilla that featured heavily in an anole character displacement versus taxon cycle debate in the early 90s.

By analyzing the size distribution of anole dentaries dating back 10,000 years on Anguilla, Melissa argues that not only is there a lack of evidence for a taxon-cycle involving A. pogus, but that there’s no evidence that A. pogus has occurred (in meaningful numbers) on the island at all! To further test this, Melissa has sequenced a portion of cytochrome b for five specimens thus far, all of which have turned out to be A. gingivinus, with more sequences coming down the pipeline.

Anoles aren’t the only lizards Melissa has uncovered – with Thecadactylus, Ameiva and Leiocephalus all present. Anoles have dominated the fauna in all but the earliest (and sparsely sampled) time slice and Leiocephalus was historically present (but probably not abundant) but disappeared several thousand of years ago and has not reappeared in the sub-fossil record since.

This was my first exposure to sub-fossil analysis of lizard communities and I was definitely impressed. I don’t have a sense of how much similar work there is in the literature (but I’m sure readers of the blog can educate me), but there seems substantial scope to collaborate with zoo-archaeologists to get more specimens and data on past trait variation and anole assemblage composition.

Where Are The Anoles?

By Adam Algar

On January 11, 2013

In All Posts

Missing anoles? Or missing anole biogeographers?

As you may (or may not) know, the 6th meeting of the International Biogeography Society (IBS) is underway in Miami, FL.

So here I am, in the anole invasion capital of the world, with the Caribbean on the doorstep, and a look through the conference program reveals a paltry TWO talks on our favourite lizards. Contrast this with Martha Munoz’s reports from SICB, with 18 (or so) anole talks. Of course, there’s something to be said for quality over quantity and the two anole presentations here do a good job of flying the flag. Yesterday Jonathan Losos kicked off one of the symposia with a talk on anole traits, function and biogeography (with a smashing blue / yellow colour scheme), and later today I’ll be searching out a poster on anole fossil assemblages (post forthcoming)

Nonetheless, I still can’t help wonder, where are the anole biogeographers? You can’t look through a text on island biogeography, species area curves or adaptive radiation without finding a dewlap. So the question is, are people not doing anole biogeography anymore? This isn’t the case, so it can only be that either the IBS isn’t on the radar, or people choose not to go. Either way, you’re missing out on a great meeting – it’s an excellent opportunity for the anole research community to radiate and reach a slightly different audience.