<img src="https://www.anoleannals.org/wp-content/uploads/2020/05/Anole-Annals-Logo-375w.png"> Anole Annals

Studies of adaptive radiation often focus on two main axes of divergence: the structural niche (e.g., where a species lives) and resource niche (e.g., what a species eats). In his SSE Symposium talk titled “The physiology of adaptive radiation,” Alex Gunderson explained the importance of a third, under-appreciated axis of species diversification: the thermal niche. Gunderson and colleagues tested whether different approaches to estimate the rates of evolution of the thermal niche lead to different conclusions, and whether thermal traits evolve at similar rates to classic ecomorphological traits like body size and limb length.

Scientists generally use three main approaches to quantify the thermal niche and estimate rates of thermal niche evolution: ecological niche modeling (ENM), organismal body temperatures, and physiological data (tolerance/sensitivity to different temperatures). Different studies use different approaches, but few use all three. Each of these metrics addresses a different scale of thermal biology, from broad environmental variables (ENM) to individual organisms (physiology). Gunderson and colleagues therefore predicted that estimated rates of evolution would vary based on the metrics used, and they used data from a number of Anolis species to test this prediction.

Specifically, the authors predicted that: a) ecological niche modeling approaches would estimate greater rates of thermal niche evolution, because environmental factors like temperature and precipitation used in ENM are very broad metrics, and are not necessarily directly correlated with individual thermal niche; b) organismal temperature data would estimate intermediate rates of thermal niche evolution, while it is a measure of individual thermal niche, it is also quite plastic; c) physiological measures would estimate the most conservative/low rates of evolution, because measures of thermal maxima and minima most accurately reflect the possible tolerance and sensitivity of individuals to thermal environments. They found that physiological data does indeed produce the most conservative estimates of thermal trait evolution, but their predictions about the performance of ENM and body temperature differed. Estimates of thermal niche evolution were highest when using body temperature data, and were intermediate when based on ENM. The fact that body temperature-based estimates of evolution rates were higher than ENM-based estimates suggests that researchers are generally underestimating error in body temperature measurements in the field.

After evaluating the results of these three different approaches in relation to thermal niche evolution, the researchers then compared rates of evolution of thermal traits to those of classical ecomorphological traits. When they used ENM, thermal traits seemed to evolve much more rapidly than morphological traits. In contrast, when they used physiological data, they found the opposite. Clearly, different metrics of climatic niche lead to different conclusions about evolutionary patterns. Gunderson therefore recommends incorporating aspects of multiple ecological and physiological scales when studying divergence of the thermal niche.

Evolution 2016: Genomic Insights into Anolis carolinensis Phylogeography

By kmwinchell

Anoles, in particular Anolis carolinensis, have long been considered an ideal group for studies investigating thermal physiology, reproductive endocrinology, and even regeneration. With the recent publication of the A. carolinensis genome (see AA posts on this here and here), the possibilities for new genomic studies in this new model species have significantly increased.

Joseph Manthey and co-authors used this new resource to clarify the phylogeographic relationships of A. carolinensis. Previous research on the phylogeography of A. carolinensis using both mitochondrial DNA and nuclear DNA showed that there were 5 clades. However, the relationships between these groups differed between the two approaches. Joseph looked at the genomes of 42 individuals from 26 localities across the native range to determine the true evolutionary relationship between regional groups and to shed light on the demographic histories of the groups. Manthey sequenced 500 loci using an anchored hybrid enrichment approach.

STRUCTURE analysis showed that the clusters had little admixture

Manthey et al. found that the genomic data predicted 5 genetic groups, in agreement with both the nuclear and mitochondrial analyses previously done. Their results also indicated that the 5 genetic clusters were distinct with little admixture. However, the relationships between groups did not agree with either the mitochondrial or nuclear trees, yet all nodes had extremely high support (93-100%)

Finally, Manthey commented on the likely timing of this diversification and associated demographic trends. Their results indicate that the initial split occurred during the late Miocene or early Pliocene and that the remaining diversification occurred during the Pleistocene. They also found that the most Southern population had a significant number of fixed genes while other populations did not. This suggests that this group was likely the oldest and most stable and supports an “out of Florida” hypothesis of diversification.

Evolution 2016: “Lightning” Strikes Three Times on Anoles in Afternoon Session: Character Displacement, Performance Trade-Offs and Opsin Evolution Matching Dewlap Color in Anoles

By mjohnson216

In this afternoon’s round of lightning talks, anoles were the focus of three fantastic (but short!) presentations on adaptation. It’s not easy to summarize a whole project in five minutes, but that’s just what these three speakers did, and each left me wanting to know more!

First, James Boyko, a Masters student working with Luke Mahler at the University of Toronto, described his work on morphological evolution in Lesser Antillean anoles. When similar species compete over a shared resource, there are two possible outcomes: extinction or divergence (i.e., character displacement). Lesser Antillean anoles are an excellent system in which to study the role of character displacement, as these islands all have either one medium-sized species, or one large and one small species. Further, the species on these islands represent two colonization events – one from the north, and one from the south. James first confirmed the classic pattern on body size evolution, finding that a three peak Ornstein-Uhlenbeck model (i.e., one that predicted large and small lizards on two-species islands, and medium lizards on single-species islands) best fit the observed data (consistent with Butler and King 2004). But when he analyzed 20 other ecologically-significant morphological traits, this three peak model did not predict trait evolution better than a model based on random chance, although the northern and southern clades significantly differed in these morphologies. In summary, to understand the evolution of Lesser Antillean anoles: body size matters, as evolution in body size is clearly an important factor to reduce inter-species competition, but lineage matters too, as body shape was predicted by ancestry.

Next came Ann Cespedes, a Ph.D. student with Simon Lailvaux at the University of New Orleans. Ann is studying functional trade-offs in green anoles (Anolis carolinensis), focusing on relationships between fitness and performance. Many studies have searched for these trade-offs in the past, and some have found them, but others haven’t. Why the discrepancies? Ann proposed that previous studies haven’t always considered sex differences in functional trade-offs, that measuring only two traits (one associated with fitness and one with performance) may not reveal real trade-offs, and that differences in individual quality are often ignored. To consider all of these factors, she measured a suite of performance and morphological traits in 60 male and 60 female green anoles. Illustrating the limitations of examining raw data on sprint speed and endurance, Ann found no suggestion of the predicted trade-off between these traits. But when using a composite measure of all performance measures (sprint speed, bite force, clinging ability, exertion, endurance, jumping ability, and climbing ability) as a control for individual quality, the trade-off between speed and endurance became clear. Males and females also differed in their speed-endurance trade-offs, as body size predicted performance in different traits between the sexes, and body shape predicted male but not female performance. So performance trade-offs do exist, but you have to know how to look for them!

To conclude the session, Alexander Stubbs, a graduate student in Jimmy McGuire’s lab at the University of California, Berkeley, described the differences between opsin gene expression in two Cayman Island anoles: Anolis sagrei (a species with a red dewlap that reflects long wavelength radiation) and Anolis conspersus (a species with a blue dewlap that reflects short wavelength radiation). Alexander proposed that these different dewlap colors might provide different selective pressures on opsins in the two species to allow better color discrimination and angular resolution. Using RNAseq to measure mRNA in the eyes of six males of each species collected at solar noon or at sunset, the results were exciting. As predicted, Anolis conspersus had higher expression of opsins that increase visual sensitivity to UV, blue, and green wavelengths, and Anolis sagrei had higher expression of opsins that increase long wavelength sensitivity. Alexander also found that gene expression different substantially between noon and sunset, and further, there was surprisingly little variation in opsin expression between lizards, in stark contrast to the wildly varying opsin expression observed in humans.

Evolution 2016: It’s Getting Cold in Here!

By kmwinchell

Tamara Fetters with her poster at Evolution 2016

Tamara Fetters, from the McGlothlin lab at Virginia Tech, reported on her ongoing work on thermal physiology in Anolis sagrei during the first poster session here at Evolution 2016 in Austin, Texas. Tamara looked at thermal tolerance and sprinting abilities at different temperatures and how that related to the latitude of the population. Specifically, she asked if lower temperatures regularly experienced by the Northern populations influence cold tolerance and performance at those temperatures. She expected that Anolis sagrei, native to Cuba and the Bahamas and introduced into the Southern U.S., would show signs of adaptation to its new, colder home in the more Northern mainland populations compared to the native range island populations in the South.

Tamara’s poster focused on two main experiments. In the first she calculated thermal tolerance to cold temperatures using a classic critical thermal minimum (CTmin) setup: with an ice bath she slowly lowered the body temperature of each animal until it was unable to right itself. This method approximates the minimum temperatures that the animals can handle in the wild. She found a clear trend showing a decrease in the minimum temperature tolerated as latitude increased. In short, Northern populations could handle the cold and Southern populations could not.

In the second experiment, Tamara acclimated the lizards to 6 temperatures ranging from 12-41 °C before running them up a track to calculate sprint speed. Tamara used an impressive 25-50 animals from each of 5 populations! She calculated sprint speed from the high-speed video she took using the program Kinovea. Tamara found that across all temperatures the most Southern population ran the slowest while the most Northern population ran the fastest, with the differences remaining fairly constant.

So what’s next for Tamara? She is planning on rearing animals in a common garden setup with some animals in hot temperatures with low variability between day and night (mimicking the native range, Southern habitats) and some animals in cool temperatures with high variability between day and night (as is experienced in the Northern habitats). She hopes that these studies will help her understand the genetic basis of this thermal tolerance and the extent of plasticity in thermal adaptation.

One last note – Tamara wanted to thank Anole Annals for helping her determine her study locations. She was able to determine which areas were likely to have Anolis sagrei and how far North they have spread because of Anole Annals posts (like this one) and comments.

Click to view a bigger version of Tamara’s poster

Evolution 2016: A Peculiar Case of Mitochondrial DNA Introgression in Puerto Rican Grass-Bush Anoles

By Jonathan Losos

On June 18, 2016

In Anole Photographs, Classics from the Literature, Introduced Anoles, Natural History Observations, Notes from the Field

FullSizeRender1

Tereza Jezkova helped kick off the anole festivities at Evolution 2016 with her talk entitled: “A peculiar case of hybridization with advantageous mtDNA introgression and lack of nuclear introgression in Caribbean anoles.” Along with a string of co-authors (Todd Castoe; Manuel Leal; Daren Card; Drew Schield; David Elzinga; Javier Rodríguez-Robles), Tereza has discovered that completely normal looking Anolis pulchellus populations in western Puerto Rico (and a bit elsewhere) harbor the DNA of the closely related A. krugi.

FullSizeRender What’s going on? Detailed examination revealed two interesting findings. First, this appears to be the result not of a single hybridization event, but minimally of 15 such events, all of them apparently quite recent. The krugi mtDNA has completely displaced the pulchellus mtDNA in these populations, and population genetic analyses rule out genetic drift as the cause. Puzzlingly, genomic analyses find absolutely no krugi nuclear DNA in these populations. The mtDNA is getting in, but not the nuclear genes. Natural selection must be at work, but how? Tereza suggested some sort of genetic mechanism that excludes the nuclear DNA of the introgressing species, somehow kicking it out, likening it to a phenomenon reported in frogs and some insects, but not in any amniotes.

Dewlap Displays in Cuban Knight Anoles (A. equestris)

By James T. Stroud

On June 17, 2016

While exploring the grounds of Fairchild Tropical Botanical Gardens with Janson Jones this past weekend, we extremely fortunately happened upon a large adult male Cuban knight anole (A. equestris) in full displaying swing. Despite the fact that knight anoles have an impressively large dewlap, I have often found this to be a relatively rare event, as large crown-giant species tend to display less than other smaller and more active species. This individual was displaying at a height of ~15 m, just below the fronds of a large Royal Palm (Roystonea regia). We didn’t see any other neighboring knight anoles, so were unsure if this was a directed or passive display series. In all, this lizard performed perhaps 4-5 sets of dewlap displays (each comprising of 4-5 dewlap extensions) before stopping and retreating back into the canopy.

Anoles typically follow a predictable and repeated pattern of display that gradually increases in intensity. Initially, and rather lethargically, an individual will nonchalantly raise its head and extend its dewlap without much extra effort (stage a); described below from Losos (2009).

Adapted from Losos (2009), which itself is adapted from Losos (1985). Aggressive behavior of A. marconoi showing three stages of increasing display intensity – note stage (c) include full body elevation alongside simultaneous tail and dewlap extensions.

This then escalates to include a slight body raise (stage b).

And ultimately results in a dramatic finale – in full display all limbs will be extended to raise both their body from the substrate (in this case the trunk of a palm tree) and elevate their tail (stage c). In the following picture you can see this final stage of displaying where intensity peaks – albeit in this individual with a regenerated (and rather stubby) tail.

Help Identify Ecuadorian Anole

By Mike Friedwald

On June 16, 2016

Hello,

A friend sent me some photos of this female anole he found in Limón Indanza, in the Morona-Santiago Province of Ecuador.

Any ideas on a possible species? I know it is not as easy as with a photo of a male.

Convergent Evolution between Pythons and Boas

By Damien Esquerre

On June 15, 2016

boa convergence

It’s not anoles but at least it’s about convergent evolution!

A recent study by Damien Esquerre and Scott Keogh published in Ecology Letters found that pythons and boas, the two famous constrictor snake families, have evolved convergent head shapes. The study was based on over 1,000 specimens and including most of the species. Pythons and boas that occupy the same micro-habitat or ecology (i.e. arboreal, terrestrial, semi-aquatic, semi-fossorial) look more like each other than to other snakes in their own family. This is exciting because it highlights how important ecology and adaptation is in shaping biological diversity.

Abstract:

Pythons and boas are globally distributed and distantly related radiations with remarkable phenotypic and ecological diversity. We tested whether pythons, boas and their relatives have evolved convergent phenotypes when they display similar ecology. We collected geometric morphometric data on head shape for 1073 specimens representing over 80% of species. We show that these two groups display strong and widespread convergence when they occupy equivalent ecological niches and that the history of phenotypic evolution strongly matches the history of ecological diversification, suggesting that both processes are strongly coupled. These results are consistent with replicated adaptive radiation in both groups. We argue that strong selective pressures related to habitat-use have driven this convergence. Pythons and boas provide a new model system for the study of macro-evolutionary patterns of morphological and ecological evolution and they do so at a deeper level of divergence and global scale than any well-established adaptive radiation model systems.

Reference:

Esquerré, D & J S Keogh. 2016. Parallel selective pressures drive convergent diversification of phenotypes in pythons and boas. Ecology Letters, 19(7): 800-809.

Predation, Food Abundance and Male-male Fighting Drives Natural Variation in Lizard Tail Autotomy

By Chi-Yun Kuo

On June 14, 2016