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Brown Anoles Invade Europe!

Reporting from Germany where she was leading a course on Transposable Elements, Jessica Stapley  — of mainland anole fame — posted this picture to Twitter. It appears brown anoles (A. sagrei) have set up a new home in one of the greenhouses at Berlin’s Botanical Gardens!

There are reports of un-established populations of green anoles (A. carolinensis) in southern Spain and the Canary Islands (reviewed here), as well as a report of Cuban knight anoles (A. equestris) also on the Canary Islands.

Does anyone know of other European records of anole populations?

 

An Experimental Test of Whether Dewlaps Are Adapted to Increase Detectability

Anolis krugi. Photo by Manuel Leal from the Leal Lab webpage

Anole biologists are fascinated by the variation in dewlap colors and patterns both within and between species. One popular hypothesis is that dewlaps are adapted to be easily detectable against the background in which they are found. A variety of tests have been published, correlating dewlap colors with ambient light, background vegetation and other characteristics. Now Alex Gunderson and colleagues have developed an experimental method of directly testing the hypothesis. Manuel Leal, a coauthor on the paper, reports on the pages of Chipojo Lab, reprinted here:

Brighter is not always better

Those that follow the Chipojoblog are familiar with one of our core tenets: strive as best you can to design experiments under natural conditions. This philosophy reflects my own view that behavior should be studied in the field whenever possible. Our recent paper in Current Zoology, “Visual playback of colorful signals in the field supports sensory drive for signal detectability,” is a prime of example of the power of this approach, in which an intimate understanding of the ecology and behavior of anoles was used to test a major prediction of the sensory drive hypothesis: are signals locally adapted? In other words, are dewlaps locally adapted to effectively grab the attention of an inattentive receiver?

Over the years we have published a series of papers supporting the hypothesis that dewlap diversity can be partially explained by selection to increase the probability of detection. However, until this paper, experimental evidence from the field was missing, in part because manipulating dewlaps of live anoles is not trivial. Furthermore, even if we were able to successfully manipulate dewlaps, there are still many other signals (e.g., body color, motion pattern, size and posture) that would be out of our control. This problem was solved by researchers working with acoustic signals a long time ago by figuring out ways to play the signal of interest in isolation in what have become known as ‘playback experiments.’ We stole a page from their book and constructed a remote-control dewlap apparatus, which provided an opportunity to display only the dewlap under natural conditions (see gizmo below).  Alex’s building and painting skills was key to the success of this gizmo. He was able to construct dewlaps with similar reflective and transmission properties of real dewlaps while taking into account the visual system of the anoles (please see papers for details).

Control-remote dewlap display apparatus. A) Acrylic box within which electrical components were housed. B) Electrical components. C) The apparatus at a mesic site with a fake dewlap displayed.

Besides presenting the dewlaps in the field, we wanted to test the hypothesis that the dewlaps are locally adapted. Under this hypothesis, increased detection in one habitat comes at the cost of decreased detection in another habitat. This functional approach to test for adaptive value of a trait is commonly used as robust evidence to support selection favoring the evolution of the trait in question. In this paper we tested if the observed differences in dewlap brightness between xeric and mesic populations of Anolis cristatellus is adaptive. If so, dewlaps from mesic populations should be more detectable in mesic habitats and dewlaps from xeric habitats should be more detectable in xeric habitats. Furthermore, detection probability should decrease in the ‘wrong’ habitat. Below are the results of the experiments.  In A. cristatellus individuals from xeric habitats have dewlaps which are darker, that is less brighter,  than individuals from mesic populations.

Responses of free-ranging A. cristatellus to fake dewlaps that mimic the brightness properties of real dewlaps.

Our findings support the sensory drive hypothesis and strongly suggest that the brightness  properties of A. cristatellus dewlaps are locally adapted via selection on signal detectability.  Furthermore, we have demonstrated that a brighter signal is not always the most detectable or effective signal. A common misconception, which is partially the result of not including the sensory system and habitat conditions as part of the analysis. Studies addressing potential functions and selective forces promoting the diversity of dewlaps found in anoles have flourished over the last decade,  nevertheless, these results are the best experimental evidence that we have to support the hypothesis that diversity of dewlap colors might be partially explained by local adaptions to habitat light conditions and the best smoking gun to support the idea that diversity of dewlap colors can be the result of local adaptations to habitat light conditions.  Additionally, our study once again underlines the need to measure both reflection and transmission when asking questions regarding the potential function of the dewlap because the two combine to determine dewlap coloration (brightness, coloration, etc.) in the real world.

Google Loves Anoles!

Ever been tempted to buy a Google Pixel cellphone? Well now you might have extra incentive! 

 

To highlight how a Google Pixel may lead you on an adventure, Google highlight’s its new photo identification feature — Google Lens — with a picture of a brown anole!

Now…how do we convince Google to give us all brand new cellphones sponsor us…?

 

HerpHighlights Podcast: an Assortment of Anoles

HerpHighlights is a Podcast run by Tom Major and Ben Marshall in which they discuss recent advances and interesting news on reptile and amphibian behavior, ecology, and conservation.

This podcast is now live and you can listen to it by clicking the link below:

https://herphighlights.podbean.com/e/026-assortment-of-anoles/

In this episode, Tom and Ben discuss many interesting research topics – both new and old – involving anoles. Notably touching on Kamath & Losos’ recent commentary on the mating systems of brown anoles (A. sagrei) in Florida, as well as Medina et al.’s review of the evolution of dorsal patterning across Caribbean anoles.

Check it out!

Kamath, A, and JB Losos. 2018. “Estimating Encounter Rates as the First Step of Sexual Selection in the Lizard Anolis Sagrei.” Proceedings of the Royal Society B: Biological Sciences 285 (1873): 20172244.

Medina, I, JB Losos, and DL Mahler. 2016. “Evolution of Dorsal Pattern Variation in Greater Antillean Anolis Lizards.” Biological Journal of the Linnean Society 120 (2): 427–35.

 

Anole Dewlapping at Bird? Part of the Fauna “Visiting” Camera Traps

Camera traps have been placed in front of some karst solution holes in the dry-transition forest of the southern slopes of the Sierra de Bahoruco, in the Pedernales province, Dominican Republic.

The holes hold water after rains, making it one of the very few spots where water is available in that forest (no surface rivers or marshes around). These holes are visited by a variety of animals, which include herps like whiptails, curlytails, racers, iguanas, and other rarities like the solenodon, and many birds.

I wanted to share this photo, since in it is the first anole recorded, an Anolis strahmi, which seems to be displaying at a nearby bird (Ovenbird, placed at extreme right of the frame).

Insular Lizards and Urban Lizards: How Brave Are They When it Comes To Foraging?

This particular lizard did not show much fear towards our plastic snake. Photo credit: Bjorn Briesen

And now for something completely different … lacertid lizards. It almost feels like I am releasing an invasive species within Anole Annals, but dr. Jonathan Losos nevertheless kindly asked me to write something about our recent paper on the effect of insularity and urbanization on the foraging decisions of the Dalmatian Wall lizard (Podarcis melisellensis), stating that “Lacertids are honorary Anoles.”

The Functional Morphology Lab of the University of Antwerp has a long-standing tradition of research on the Croatian islands. Once a year, several members of our team travel to Croatia, accompanied by a bunch of students in the context of a course on Island Biology. In fact, four out of six collaborators on our paper (including me) were still undergraduate students when conducting the fieldwork. We soon came up with the idea to compare risk-taking and reaction to novelty of wall lizards during foraging in both urban and insular environments. Strange choice? Not at all. Both urban and insular habitats are characterized by reduced levels of predation, but also a lack of resources (e.g. lower abundance and diversity of arthropod prey). Such conditions should prompt both insular and urban lizards to behave bolder during foraging, and act less neophobic, since the risk of predation is low, but the value of the food reward is high. Fortunately, the Croatian islands are perfect to test these kind of hypotheses. Sampling both rural and urban areas on the larger island of Vis was a piece of cake, especially since wall lizards can easily be found in the harbour of Vis.

Believe it or not, wall lizards can easily be found in the harbour of vis. Photo credit: Gilles De Meester

Vis itself is surrounded by many small islets, most of them inhabited by dense populations of lizards. Some of these islets, like Biševo and Veli Budikovac, are (semi-)inhabited. Others, like Brusnik, are completely devoid of any human activity. So, small islets? Check. Urban and rural areas? Check. What else was needed? A predator model to test risk-taking, and a novel object to test neophobia. After a few pilot trails in which we compared the response of lizards to a plastic snake model and a stuffed rook (Corvus frugilegus), it was clear that lizards showed more fear of the snake. Our novel object was randomly chosen to be a flashy red drinking bottle.

Lizards from smaller islets did indeed take more risks while foraging compared to lizards from the main island. Islet lizards were more likely to leave a safe refuge in order to get a tasty mealworm, despite the presence of the snake model, and also on average left their refuges faster to do so. In fact, on multiple occasions, lizards did completely ignore the snake model, even running over it to get to the petri dish with food.

Another example of a very bold lizard. Photo credit: Bjorn Briesen.

But even on the main island of Vis, lizards behaved very boldly. In fact, between observations, we were often capable of hand-feeding some wild individuals. On the island of Brusnik, a few lizards showed absolutely no fear at all of humans, even licking and biting our hands when we came close to remove the petri dish at the end of a trail. This is, of course, a very nice illustration of  the phenomenon known as “island tameness.” Due to a lower number of predators on islands and islets, insular species often lack normal antipredator behaviour, something that even Darwin already reported after his famous voyage on the Beagle. Unfortunately, this tameness is the main reason for the historical decline and extinction of many insular species, especially after humans started bringing new predators (rats and cats) to these islands.

Maybe he was just thirsty? Photo credit: Gilles De Meester

Neophobia, whether the lizards were willing to feed in front of a novel object, did not differ between main island and small islets, despite the assumption that islet lizards should behave less neophobically as they will benefit more from exploiting new resources. In fairness, lizards overall seemed to lack a neophobic reaction towards the novel object, as they did not differ in feeding latencies from the control group. Interestingly, however, in Brusnik, lizards behaved very neophilically towards the drinking bottle. Some individuals even completely ignored the offered food in favour of this novel object.

Surprisingly, lizard behaviour did not differ between urban and rural environments. While numerous papers have proven that urban birds and mammals  behave more boldly and less neophobically compared to their rural conspecifics, the few studies on reptiles have yielded mixed results. In fact, it is still unclear whether lizards do indeed experience lower predation risk in urban areas. While natural predators might be rare, they are often replaced by dogs, rats or cats (and we all know that cats are the killing machines of nature). We did indeed observe predation on wall lizards in the harbour of Vis, not by a cat, but  by a hooded crow (Corvus cornix). A strong argument can be made that urban habitats are more open, leaving foraging lizards at greater risk of being seen by predators.

More detailed studies of behaviour of urban lizards, together with estimations of resource availability and predation pressure, could help us understand why lizards seemingly adapt different to urban environments compared to mammals or birds.

More detailed interpretation of our results, complete with nice graphs and interesting p-values, can be found in our published paper.

Variation in Cold Tolerance across the Geographic Range of the Green Anole: How Do More Northern Populations Withstand Colder Temperatures?

From Card et al. 2018

Shane Campbell-Staton and colleagues have just published a paper in Molecular Ecology on the physiological and regulatory basis of variation in cold tolerance across the range of Anolis carolinensis. In the same issue, Daren Card and colleagues have written a very nice, freely available, summary of that article. Here’s the abstract from Card et al.’s review:

How does climate variation limit the range of species and what does it take for species to colonize new regions? In this issue of Molecular Ecology, Campbell-Staton et al. (2018) address these broad questions by investigating cold tolerance adaptation in the green anole lizard (Anolis carolinensis) across a latitudinal transect. By integrating physiological data, gene expression data and acclimation experiments, the authors disentangle the mechanisms underlying cold adaptation. They first establish that cold tolerance adaptation in Anolis lizards follows the predictions of the oxygen- and capacity-limited thermal tolerance hypothesis, which states that organisms are limited by temperature thresholds at which oxygen supply cannot meet demand. They then explore the drivers of cold tolerance at a finer scale, finding evidence that northern populations are adapted to cooler thermal regimes and that both phenotypic plasticity and heritable genetic variation contribute to cold tolerance. The integration of physiological and gene expression data further highlights the varied mechanisms that drive cold tolerance adaptation in Anolis lizards, including both supply-side and demand-side adaptations that improve oxygen economy. Altogether, their work provides new insight into the physiological and genetic mechanisms underlying adaptation to new climatic niches and demonstrates that cold tolerance in northern lizard populations is achieved through the synergy of physiological plasticity and local genetic adaptation for thermal performance.

Do Dead Green Anoles Turn Blue?

Photo by Dee Simpson

AA reader Dee Simpson reports:

I recently found a deceased Green (Carolina) Anole near my home in central Florida. What struck me is that it was blue. At first, I thought just looked blue because it was desiccated, but on further examining the picture, I realized that one leg was green – if it was just the decaying process, I would expect the whole thing to be the same color/state. I came across the entry on your Anole Annals page regarding blue Carolina Anoles in Florida and was wonder if this could be one of those? Or is it just at a stage of decomposition where the color is weird?

Another Three-Legged Anole…but with a Plot Twist

Anyone who has spent a considerable amount of time catching anoles in the field has seen their fair share of injured animals. Many species we commonly study (e.g. brown anoles) are just the perfect size to be a snack for any hungry predator (and even humans! see this). Several previous posts have documented adult anoles that have sustained severe injuries (limb loss – see my previous post) and survived. But can these animals thrive with such injuries or do they just limp along through life?

Here I add to this string of anecdotes with a unique datum. This female Puerto Rican crested anole was caught by none other than James Stroud and Chris Thawley at Fairchild Botanical Gardens just this week. She is missing the rear right foot (not an unusual injury). What is new here is that I dissected this female as part of a study conducted by James, Chris, and myself, and I can report that this female, despite her handicap, is not only alive but seems to be thriving. Compared to a cohort of females captured at the same time and place (n= 13), she has greater body condition and fat mass than most of her cohort (Figure 1) and is reproductive at stage 4 (Gorman & Licht 1974). For those unfamiliar, stage 4 means that she has two developing eggs (1 in each oviduct). The mean stage for the cohort is 2.92, and, thus, her reproductive stage is more advanced than the majority of the cohort (only 2 of 13 individuals at stage 4).

Figure 1. Fat mass and residual body condition (log(mass) x log(SVL)) for the entire cohort (gray circles) and the injured female (black circle). There was no relationship between SVL and fat mass for females in this cohort so fat mass is not corrected for body size.

Cox and Calsbeek (2010) demonstrated that gravid anoles have reduced locomotor performance and lower survival than non-reproductive females. However, this female, despite the use of only 3 good limbs, has clearly been able to procure sufficient resources to  fuel reproduction and retain a level of fat reserves above most individuals in her population.  For this reason, we denote her ‘supermom’ and concede the possibility that missing a foot or limb may not severely reduce fitness for some individuals.

Cox, R.M. and Calsbeek, R., 2010. SEVERE COSTS OF REPRODUCTION PERSIST IN ANOLIS LIZARDS DESPITE THE EVOLUTION OF A SINGLE‐EGG CLUTCH. Evolution64(5), pp.1321-1330.

Gorman, G.C. and Licht, P., 1974. Seasonality in ovarian cycles among tropical Anolis lizards. Ecology55(2), pp.360-369.

 

Replicated Physiological Diversification in Lizard Adaptive Radiation

Anolis gundlachi prefers to perch in the shade near the ground — according to a new study, these microclimate differences drove some of the later diversification of anoles.
Credit: Manueal Leal, MU Division of Biological Sciences
 Report of a new study from the webpages of ScienceDaily, produced initially by the University of Missouri:

The islands of Puerto Rico, Cuba, Hispaniola, and Jamaica — collectively known as the Greater Antilles — are home to more than 100 species of Anolis lizards. The success of this colorful group of reptiles is often attributed to the evolution of distinct body shapes and behaviors that allow species to occupy different ecological niches. A new study from an international team of biologists including from the University of Missouri reports that the evolution of physiological differences that allow these lizards to take advantage of different microclimates (e.g., sun vs. shade) may have been just as important as these physical differences. The study, which was published recently in The Proceedings of the Royal Society B, has implications for predicting how well these lizards will cope with climate change.

“Why are there so many species of anoles? That’s the big question,” says Manuel Leal, an associate professor of biological sciences at MU and one of the authors of the report. “The notion that morphological differences alone drove the amazing diversity of anoles is missing an important part of the puzzle.”

For scientists, the Greater Antillean anoles represent a classic example of an evolutionary process known as adaptive radiation. After appearing on each of the four islands about 50 million years ago, the colorful lizards quickly diversified to exploit different niches on the island’s trees, including the canopy, trunk near the ground, mid-trunk, and other twigs. Each new species developed its own distinct body type, called an ecomorph, adapted to the niche where it lived. According to Leal, this focus on differences in appearance leaves some important questions unanswered.

“How can similar species coexist without outcompeting one another? One of the tenants of evolutionary ecology is that when a structural niche is filled, species diversification should either slow or come to an end due to competition. There must be some other way they are sharing that habitat to avoid competition,” he said.

The researchers hypothesized that the evolution of physiological traits related to temperature tolerance also facilitated the maintenance of biological diversity by providing an additional axis of co-existence.

Working with Alex Gunderson with the University of California at Berkeley and D. Luke Mahler with the University of Toronto, Leal set out to test this hypothesis. The team caught and collected thermal physiological data on over 300 anoles. Most of the anoles belonged to the Puerto Rican cristatellus group, which includes four pairs of sister species, each of which occupies a different thermal niche. They also included data on Jamaican anoles. The researchers measured two aspects of thermal physiology: maximum thermal tolerance and optimal temperature for sprint performance, which they used as a measure of fitness. They asked if a species heat tolerance correlated with its optimal sprint performance. They expected that sister species would diverge in one or both of these physiological traits.

They found that all Puerto Rican species pairs diverged in at least one of the two physiological traits. In three of the four pairs, the species that preferred the warmer environment had a higher thermal maximum temperature. In two cases, the species that preferred the warmer environment also had had a higher optimal temperature. They found a similar pattern among the Jamaican anoles.

“These findings show that when morphologically similar species co-occur in Puerto Rico and Jamaica, they differ in thermal physiology. We can say that thermal physiological differentiation is important for increasing local species richness,” said Leal.

An additional insight was that thermal physiology evolved slower than morphology. This evolutionary interplay, Leal said, has real-world implications when one considers the rate at which the world’s climate is warming.

“This is not good news for the ability of anoles to adapt to climate change,” said Leal. “The data suggest that the rate at which physiology changes in anoles is not fast enough to cope with how fast temperatures are rising.”

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