Category Archives: New Research

The Lonely Clouded Anole on a Pacific Island

Anolis nebulosus

Anolis nebulosus. Photo by Hugo Siliceo-Cantero.

By H. Hugo Siliceo-Cantero and A. Garcia

In the late 1980´s, the scientists Bradford C. Lister and Andrés García discovered an interesting population of clouded anoles inhabiting the small 3.3 ha island of San Agustin located just off the Pacific coast of Jalisco, Mexico. This island was also close to the actual protected area of tropical dry forest on the mainland in the Chamela-Cuixmala Biosphere Reserve. Lister and García reported that the abundant anole population on San Agustin was maintained a decade later at much higher densities than the mainland population. We began to study this population in 2007 as a graduate student. Since then, we have studied several aspects of the ecology of this island population comparing this with the ecology of anoles on the mainland.

The existence of such island populations enables scientists to carry out natural experiments that provide invaluable information helping us to understand ecological and evolutionary processes.

This Clouded Anole (Anolis nebulosus) species that is on San Agustin Island is endemic to Mexico, and is of particular interest as this population has evolved in the absence of similar species of the same genus, or congeners. The species on the island also occupies a broad niche of perch height and a low number of lamellae, and is one of the most sedentary anoles known. Our work demonstrated that San Agustin population of the Clouded Anole has distinct morphological and genetic traits compared to conspecifics on the mainland.

Recently, we found that the insular population also presents distinct ecologic traits compared to those of the mainland population. In our manuscript “Assessing the relative importance of intra- and interspecific interactions on the ecology of Anolis nebulosus lizards from an island vs. a mainland population”, we suggest that the processes that drives the ecology and evolution of this insular population (intraspecific competition) differs from those that are important in the mainland (interspecific competition).

We believe that the results of our research on the insular population of anoles on San Agustin Island complement the scenario of Caribbean anoles, where congeneric competition is the key evolutionary driver. Furthermore, in our study, we used video cameras to provide direct evidence of predation, interspecific and intraspecific encounters and aggression, which was possible because the Clouded Anole is a sedentary lizard.

It has been a pleasant and rewarding experience for me to study the Clouded Anole. Although spending hours in the field observing a largely sedentary lizard may seem a little boring and tedious, the data from our studies have revealed a fascinating adaptation to the natural and social environment with unique physical, genetic, and ecological characteristics.

Currently, the population of Clouded Anoles on San Agustin has been dramatically reduced, almost to the point of extirpation. We think that two natural events, the hurricanes Jova in 2011 and Patricia in 2015, as well as invasive studies such as Hernández-Salinas et al. (2016) where they extracted 77 anoles from this small island, are the cause of the dramatic reduction in the Clouded Anole of San Agustin Island. As ecologists, we believe that research should not be done at the expense of the species or population under study, but should ensure that the population remains intact to continue along its evolutionary path, and further elucidate our understanding of the natural world around us.

We are currently monitoring both insular and mainland populations in order to understand and evidence the ecological implications of such natural and anthropogenic reduction on anole populations.

Clipped Claws and Consequences for Anolis Adhesive Performance

Figure 1. Differences in claw clipping used in Bloch and Irschick (2005) and our study. (A) The entire claw was clipped after the distal end of the toe pad. (B) In our study, we partially clipped the distalmost portion of the claw.

Figure 1. Differences in claw clipping used in Bloch and Irschick (2005) and our study. (A) Bloch and Irschick (2005) clipped the entire claw after the distal end of the toe pad. (B) In our study, we partially clipped the distalmost portion of the claw.

Toe and claw clipping are common techniques used to identify individuals in mark and recapture studies, but their impacts on whole organism performance are unclear (Dunham et al., 1988). Anoles have not only developed subdigital adhesive toe pads to promote adhesion on relatively smooth substrates, but have also retained claws to enhance attachment to rough substrates (Irschick et al., 1996; Zani, 2000). Thus, clipping entire toes or claws may have drastic effects on the clinging ability of anoles or other adhesive pad-bearing lizards. In our recent article published in Acta Herpetologica, my co-authors and I investigated how partially removing the claws of brown anoles affects their adhesive performance.

Figure 2. Mean maximum clinging force of Anolis sagrei with intact and partially clipped claws. Overall, partial claw clipping had no significant effect on maximum clinging ability.

Figure 2. Mean maximum clinging force of Anolis sagrei with intact and partially clipped claws. Overall, partial claw clipping had no significant effect on maximum clinging ability.

Bloch and Irschick (2005) removed entire claws from Anolis carolinensis (Fig. 1A) and measured its impact on their clinging ability. Not surprisingly, claw removal resulted in a significant decrease in the clinging ability of A. carolinensis, likely a consequence of the severing of flexor tendons that are critical in adhesive toe pad engagement. In an effort to test this hypothesis and preserve these tendons, we used a motorized force sensor (Niewiarowski et al., 2008) to measure the maximum clinging ability of 19 Anolis sagrei before and after their claws were partially clipped (Fig. 1B).

Overall, we found that partial claw clipping did not significantly impact maximum clinging ability (Figure 2). This suggests that clipping the entire claws of anoles may indeed sever the flexor tendons crucial to toe pad engagement. Furthermore, we expected clinging ability to increase after partial claw clipping because claws should theoretically interfere with the contact the subdigital adhesive pads are capable of producing. However, this did not appear to be the case, suggesting that claws may not inhibit the engagement of subdigital pads or that morphological features and/or behavioral traits reduce the effect of this interaction.

Anolis sagrei

Anolis sagrei

Although permanent marking solutions would be most beneficial for mark and recapture studies, partial claw clipping may be a useful alternative for shorter-term studies because it does not appear to reduce adhesive performance on smooth substrates. Future work should further consider the interactions between subdigital adhesive toe pads and claws, and determine the possible ramifications for adhesion and adhesive locomotion, particularly on rough substrates. Be sure to check out our full article for more details!


Bloch, N., Irschick, D.J. (2005): Toe-clipping dramatically reduces clinging performance in a pad-bearing lizard (Anolis carolinensis). J. Herpetol. 39: 288-293.

Dunham, A.E., Morin, P.J., Wilbur, H.M. (1988): Methods for the study of reptile populations. In: Biology of the Reptilia, pp. 331-386. Gans, C. Huey, R.B., Eds, Alan R. Liss, Inc., New York.

Irschick, D.J., Austin, C.C., Petren, K., Fisher, R.N., Losos, J.B., Ellers, O. (1996): A comparative analysis of clinging ability among pad-bearing lizards. Biol. J. Linn. Soc. 59: 21-35.

Niewiarowski, P.H., Lopez, S., Ge, L., Hagan, E., Dhinojwala, A. (2008): Sticky gecko feet: the role of temperature and humidity. PLoS ONE 3: e2192.

Zani, P. (2000): The comparative evolution of lizard claw and toe morphology and clinging performance. J. Evol. Biol. 13: 316-325.


SICB 2018: When the Lights Go Up in the City

Chris Thawley at a crossroads.

Chris Thawley at a crossroads.

Plants and animals across the globe are dealing with increasingly changing environments resulting from urbanization. One such habitat alteration is artificial light at night (ALAN) that may affect how animals acquire or use energy. Because brown anoles (Anolis sagrei) are common invaders of urban habitats, they make good models to examine the consequences associated with ALAN. Thus, Chris Thawley of Jason Kolbe’s lab at University of Rhode Island altered the level of ALAN on female brown anoles to examine how ALAN might affect morphology, reproduction, and stress.

If this sounds familiar, Chris talked about this work at the 2017 JMIH meeting, which caught Anole Annals press. To recap, Chris found that ALAN increased female growth, advanced the start of egg laying to earlier in the season, and increased the reproductive output of smaller females. However, he hypothesized that such beneficial effects would be countered by negative effects on other traits. Thus, Chris measured levels of the stress hormone corticosterone in the blood of females, hypothesizing that those exposed to ALAN would have higher stress. Although marginally non-significant, females actually tended to have lower corticosterone levels. Chris presented new data for this presentation showing that male corticosterone levels were unaffected by ALAN too, suggesting neither adult male nor female brown anoles have a stress response to artificial light.

Thus, it appears ALAN exposure over this 7-week study was beneficial for brown anole reproduction. However, Chris cautioned that there may be negative consequences on other traits such as immunity or HPA function. ALAN might also induce negative consequences for reproduction later in life, such as a reduced lifespan. I recommend keeping an eye on the Kolbe lab to find out!

SICB 2018: Moms Help Embryos Beat the Heat

Putter, Austin, and a real big tree they visited while travelling to the meeting.

Putter, Austin, and a real big tree they visited while travelling to the meeting.

The effect of urbanization on animals was the topic of many presentations at this year’s SICB meeting. One difference in the abiotic environment of urban areas is that they are often hotter than neighboring natural areas. Sarin “Putter” Tiatragul and colleagues (Josh Hall, Nathaniel Palik, and Dan Warner) at Auburn University are interested in whether urban environments might influence the nesting ecology and development of anoles. Thus, they set to the field to search for nest sites of the Puerto Rican Crested Anole (Anolis cristatellus).

Putter predicted females would choose warm, open-canopied nest sites at both urban and forested habitats, but that the availability of such locations would not be equal between sites. As predicted, randomly available areas in urban habitat had less tree cover and were warmer than randomly available locations in the forest. In the forest, females nested in locations that were similar to what was randomly available (no preference) in terms of distance to the nearest tree, canopy cover, and nest temperature. However, urban anoles nested in less open areas and closer to trees than what was randomly available in the urban habitat. This resulted in female-chosen nests sites being cooler than what was randomly available.

These findings suggest female anoles in forested areas are not choosing nest sites, probably because the forested habitat is homogenous and provides little variation to choose amongst. However, females in urban areas search out cooler microhabitats possibly to achieve favorable incubation conditions for their offspring. Putter also suggested these females may be simply nesting close to where they normally occur, which is close to trees. Either way, females are using the habitat differently in urban areas and such variation will likely have consequences for offspring during development.

SICB 2018: Revisiting the Fitch-Hillis Hypothesis in Mexican Anoles

A small sample of anole dewlap diversity. Image from Nicholson et al. (2007).

A small sample of anole dewlap diversity. Image from Nicholson et al. (2007).

Dewlaps are pretty dazzling, ranging in size, coloration, and sexual dimorphism substantially among the 400+ species of anole currently recognized. Levi Gray, a doctoral candidate at the University of New Mexico is fascinated by Anolis dewlaps, and has spent many years studying them. One of the classic hypotheses surrounding dewlap evolution in anoles is that its size follows a clinal pattern with environment (Fitch and Hillis 1984). In their formulation, Henry Fitch and David Hillis proposed that, due to a relatively short breeding season, anoles in more seasonal habitats have larger dewlaps than anoles in more aseasonal habitats. This hypothesis makes an explicit connection between the intensity of sexual selection and the size of a conspicuous ornament.

Levi set out to test the Fitch-Hillis hypothesis in 40 species of Mexican anoles distributed across environmental gradients, with some species found in aseaonal cloud forests and rainforests and others found in more seasonally dry habitats. Contrary to the Fitch-Hillis hypothesis, he found no relationship between seasonality and dewlap size in the Mexican anoles. He did detect a few clade effects: for example, a group of closely related western Mexican anoles all have large dewlaps. He then examined the Fitch-Hillis hypothesis within a single widespread species of anole, A. sericeus, to see if the pattern holds up within species, even if it doesn’t hold up among species. Again, he didn’t detect a pattern. Levi suspects that the relatively limited sampling of the original study might have led to a pattern that doesn’t hold up when a broader sampling within and among species is employed. It is possible that seasonality impacts a different aspect of the dewlap, such as coloration, but this remains untested. Levi’s results suggest that the processes impacting dewlap size might be complex, and promises more to come. Stay tuned!

Levi Gray presents his research on dewlap size evolution in Mexican anoles at SICB 2018 in San Francisco.

Levi Gray presents his research on dewlap size evolution in Mexican anoles at SICB 2018 in San Francisco.

SICB 2018 – Are Anoles Adapting to Hot City Environments?

Urbanization, the creation and spread of urban habitats, is increasing across the world. Species that live in these urban habitats are subject to many alterations in their environment, including changes in food, predators, noise, and light among others. One of the most well-known changes associated with cities is the “Urban Heat Island” effect, where city habitats are hotter than surrounding areas due to increases in pavement and other heat-absorbing materials. For lizards such as anoles, living in this hotter environment could be challenging, as increased heat could reduce time available for foraging for food or defending territories, or, in more serious cases, might even lead to death. Shane Campbell-Staton, a postdoctoral researcher at the University of Illinois and the University of Montana, decided to test if anoles were adapting to these hot urban environments, and, if so, what mechanisms were driving this adaptation.


Cities are hotter than the surrounding landscape.

Shane worked with crested anoles (Anolis cristatellus) from four different areas of Puerto Rico that had both urban and nearby natural environments. He and Kristin Winchell, his coauthor, verified that anoles in these urban habitats did indeed experience hotter conditions, and that, as a result, their body temperatures were also higher than anoles from nearby natural areas. In the lab, Shane found that these city anoles were capable of tolerating higher temperatures than their counterparts from natural areas as well. However, after 8 weeks in the lab, anoles from both types of habitats had similar temperature tolerances. Shane also raised offspring from these anoles under common conditions in the lab and found that these offspring had similar temperature tolerances (thermal limits), regardless of whether they came from urban or natural environments. These results show that anoles can have a plastic response to the thermal conditions in their environment, meaning that the differences Shane and Kristin saw in Puerto Rico are induced by an anole’s exposure to temperatures and are not completely determined by their genes.

Crested anoles (Anolis cristatellus) make use of many human-altered habitats.

Crested anoles (Anolis cristatellus) make use of many human-altered habitats. Photo by Andrew Battles.

Shane, however, continued to explore this question: he wanted to know if the ability, or plasticity, of an anole to alter its thermal tolerance in response to exposure to high urban temperatures was due to changes in its genetic structure. In essence, he wanted to know if anoles had evolved a higher responsiveness (or plasticity) in response to inhabiting hotter, city habitats. To get at this, Shane exposed anoles to both hot and normal temperatures in the lab and looked at their levels of gene expression. Using a transcriptomics approach, Shane could see which genes were activated differently when lizards were exposed to temperatures indicative of city and natural habitats. Shane observed differences in variation in the genes in use at these temperatures. He also found higher levels of differentiation between genes involved in thermal adaptation between lizards from city and natural environments. These exciting results show that living in hotter city environments has selected for lizards which are more able to respond to these hot temperatures when they experience them. Shane is continuing to dig deeper into these data to determine which specific genes may have been altered to understand the mechanisms by which lizards are able to alter their heat tolerances. We’re looking forward to seeing these results at a future conference!

On a side note, Shane will be setting up his own lab at UCLA this year, and he’ll be looking for talented graduate students interested in physiology, adaptation, and genomics. Don’t hesitate to look him up!

Register Now for the 2018 Anole Symposium

Winning Symposium t-shirt design by Eric-Alain Parker

Winning Symposium t-shirt design by Eric-Alain Parker

Calling all anole aficionados! Spots for the Seventh Anole Symposium on March 17-18, 2018 at the Fairchild Tropical Garden in Miami, Florida are starting to fill up! We have a limited number of spots for attendees and fewer still for presentations. If you have already registered, great! If you haven’t, please do so soon to guarantee your spots. Registering now does not require you to also pay now  – registrants will be contacted in early 2018 to pay the $100 registration fee. Submitting an abstract is not also required at this stage, and can be amended to the registration at a later date.


The Not-So-Bitter Future of Coffee: Anolis Lizards as Biocontrol Agents in Mainland and Island Agroecosystems

Figure 7. Anolis gundlachi, Orocovis, Puerto Rico.

Figure 1. Anolis gundlachi, Orocovis, Puerto Rico.

The agroecosystems that produce the life-sustaining stimulant we know as “coffee” have long been used as model systems to study complex ecological interactions and ecosystem services, with numerous studies revealing trophic interactions among coffee plants, pests, and pest-predators. Despite the high abundance and overlapping distribution of Anolis lizards, relatively few studies have addressed their functional role in agriculture. In our recent study titled, “Anolis Lizards as Biocontrol Agents in Mainland and Island Agroecosystems,” my colleagues and I explore the biocontrol potential of anoles against the world’s most devastating coffee pest, the coffee berry borer (Coleoptera: Hypothenemus hampei) in mainland and island settings.

My vision of agricultural landscapes as post-apocalyptic biodiversity deserts was trumped the minute I stepped foot onto a shade coffee farm in Orocovis, Puerto Rico. Far from the dystopian nightmare that I had envisioned, this diversified shade coffee farm bustled with the herpetological glory and natural complexity of a native forest (Fig. 1). Furthermore – and perhaps most importantly – the farmer complained not of issues with crop yield, pests, and disease.

As a plant, coffee occurs naturally in the forest understory and is cultivated traditionally among native shade trees as an understory crop. While pressures to increase production have led many farmers to transition to more intensive practices (i.e., the reduction of shade cover and application of agrochemicals to manage crop pests), these methods are becoming increasingly unsustainable and insufficient in light of emerging biological threats. In addition to climate change and the emerging coffee rust disease, the coffee berry borer poses a unique threat for dozens of coffee growing nations and nearly 20 million small-scale farmers who depend on coffee production as a primary commodity and means of subsistence. While the coffee berry borer (CBB) is capable of inducing 60-90% reductions in yields and persists unaffected by topical pesticides, our understanding of the predator-prey interactions that drive its top-down control and how these factors vary across management regimes and eco-geographic space has profound socio-economic and environmental implications for biological control.

Representative photographs of diversified shade coffee in Mexico (a), diversified shade coffee in Puerto Rico (b), intensive sun coffee in Mexico (c), and intensive sun coffee in Puerto Rico (d).

Figure 2. Representative photographs of diversified shade coffee in Mexico (a), diversified shade coffee in Puerto Rico (b), intensive sun coffee in Mexico (c), and intensive sun coffee in Puerto Rico (d).

To assess the biocontrol capacity of anoles, we conducted experimental and field-based tests of how CBB populations respond to anole predation across mainland (Mexico) and island (Puerto Rico) coffee farms with parallel forms of land-use intensity. Anole functional response and infestation reduction potential were assessed by simulating pest outbreaks in the lab, while coffee farms were surveyed along complementary gradients of intensification. Organic, diversified shade coffee farms were representative of low-intensity production, and sun coffee monocultures that included the application of agrochemicals were representative of high intensification (Fig. 2). Continue reading The Not-So-Bitter Future of Coffee: Anolis Lizards as Biocontrol Agents in Mainland and Island Agroecosystems

Evidence for evolutionary determinism in the signal design of lizards?

Photographs of a subset of lacertid lizard species used in this study. From the left top to the right bottom: Acanthodactylus beershebensis, Lacerta bilineata, Dalmatolacerta oxycephala, Podarcis melisellensis, Tropidosaura gularis, Podarcis siculus, Heliobolus lugubris, Algyroides nigropunctatus, Lacerta media.

Photographs of a subset of lacertid lizard species used in this study. From the left top to the right bottom: Acanthodactylus beershebensis, Lacerta bilineata, Dalmatolacerta oxycephala, Podarcis melisellensis, Tropidosaura gularis, Podarcis siculus, Heliobolus lugubris, Algyroides nigropunctatus, Lacerta media.

The vast array of signals used in animal communication is a continuous source of awe and a hot topic in evolutionary and behavioral research. One important factor contributing to the signal diversity we witness today is ‘signal efficacy’: the ability of a signal to travel efficiently through the environment and attract the receiver’s attention. With this in mind, natural selection is expected to mold signal design for maximum efficacy of information transmission and detectability, leading to signal variation among populations/species living in different environments. To illustrate, a recent study by Tess Driessens and colleagues assessed the degree of variation in the dewlap design of Anolis sagrei by comparing 17 populations distributed across the Caribbean (Fig. 1).

Phylogenetic relationships among seventeen Anolis sagrei populations. Pie charts illustrate dewlap pattern proportions for each population per sex (black = solid; light grey = marginal; dark grey = spotted). Photographs represent male and female dewlaps of typical individuals from every population.

Fig. 1 — Phylogenetic relationships among 17 Anolis sagrei populations. Pie charts illustrate dewlap pattern proportions for each population per sex (black = solid; light grey = marginal; dark grey = spotted). Photographs represent male and female dewlaps of typical individuals from every population.

Their findings showed large interpopulational variation in dewlap size, pattern, and color, and more interesting, they established a link between the dewlap design of brown anoles and the environment they live in. Lizards occurring in more ‘xeric’ environments had a higher proportion of solid dewlaps with a higher UV reflectance; lizards inhabiting ‘mesic’ environments had predominantly marginal dewlaps showing high reflectance in red. This was true for both males and females. Like Ng et al. (2011) and their observations on dewlap variation in A. distichus across an environmental gradient, Driessens et al. (2017) interpret their findings as evidence for adaptive divergence of a signaling apparatus.

Surprisingly though, while there are numerous great examples of comparative studies finding support for convergent evolution in visual and acoustic signaling systems, (e.g. Endler 1992; Fleishman 1992; Nicholls & Goldizen 2006, to say a few), similar (comparative) studies, but then, on the phenotype of chemical signals are almost entirely lacking. This is probably due to the combination of only very recent developments in chemical analytical and statistical comparative tools, the time researchers need to assemble a large-scale multi-species chemical dataset, and perhaps due to our own predisposition to visual and auditory signals. Currently, the proper analytical tools for studying natural products chemistry are available and affordable, permitting comprehensive taxon-wide research on the evolution of chemical signal diversity and design. Ultimately, there has never been a better time as now to be a comparative chemical ecologist.

Photograph of the cloacal region of a male lacertid lizard (Lacerta agilis), showing his numerous femoral pores with protruding glandular secretion.

Photograph of the cloacal region of a male lacertid lizard (Lacerta agilis), showing his numerous femoral pores with protruding glandular secretion.

Finally, three Belgians, two Spaniards and one Greek (sounds like the start of a joke with ample potential) took up the challenge to examine variation in the chemical signal design of lizards. Although underrepresented in studies on chemical signal diversity, lizards are an excellent group for investigating chemical signal evolution, as many of them they bear numerous glands on their thighs that secrete waxy substances, which they deposit while moving through their habitat. These secretions are often considered the leading source of chemical signals involved in lizard communication.

The study started with a quest. A quest to collect gland secretions of as many species as possible (within a PhD timeframe). Luckily, we were fortunate enough to be able to count on the help of many collaborators (Shai Meiri, Chris Broeckhoven, …). We focussed on lacertid lizards, as they are a species-rich family distributed over a wide geographical area, and known to rely strongly on chemical communication in several contexts.

In total, we sampled secretions from 64 species throughout, Europe, Africa, and Asia, covering a wide array of habitats and climate regions: from the Mediterranean maquis over the alpine meadows in the Pyrenees Mountains, to the sandy Israeli dunes and the Kalahari Desert of South Africa (Fig. 2). Back in the lab, we determined the chemical composition and chemical ‘richness’ (number of different chemical compounds) of the secretions using GC-MS, and obtained climate data for all catch-localities from online databases.

Map showing the sample localities of the 64 lizard species under study.

Fig. 2 — Map showing the sample localities of the 64 lizard species under study.

Our gathered data showed considerable variation in the chemical richness and composition of lacertid secretion. Shared-ancestry failed to explain among-species patterns of variation, hinting that chemical signals may change relative rapidly. Most interestingly, our findings revealed a strong relationship between the environmental conditions species live in and the chemical composition of their glandular secretions. On the one hand, lizards living in ‘xeric’ environments, characterized by high temperatures and arid conditions, contained higher proportions of stable and heavy-weight compounds in their secretions. Hot and dry conditions increase the evaporation rate of chemicals, and so, decreasing the longevity of a signal. Stable and heavy-weight compounds most likely reduce evaporation rate and counteract the rapid signal fade-out through evaporation, generating a highly persistent scent-mark. On the other hand, species inhabiting wet, humid conditions produced highly aromatic and low-weight secretions containing numerous different compounds. This chemical mix probably creates a volatile-rich signal that can be used for long-distance airborne communication.

While we cannot deny that these findings of convergent evolution in the design of chemicals signals are fascinating, some would say this outcome is not unexpected.

“[…] a cadre of scientists has taken the […] view, that convergence is the expectation, that it is pervasive, and that we should not be surprised to discover that multiple species […] have evolved the same features to adapt to similar environmental circumstances. From this perceived ubiquity, the scientists draw a broader conclusion: evolution is deterministic, driven by natural selection to repeatedly evolve the same adaptive solutions to problems posed buy the environment. — J. Losos (Improbable Destinies, p. 33)

Nonetheless, I am confident to state that using by far the largest comparative dataset amassed to-date to examine patterns of chemical signal divergence, we have provided strong evidence for a significant relationship between chemical signal design and prevailing environmental conditions, which may results from differential selection on signaling efficacy (Baeckens et al. 2017).

Condition Dependence of Shared Traits Differs between Sympatric Anolis Lizards

A male slender anole (Anolis limifrons)

A male slender anole (Anolis limifrons)

A walk through a tropical rainforest can reveal astonishing forms and colors of organisms – from vibrant poison frogs and coral snakes to the vegetative camouflage of stick insects and other cryptic creatures. Perhaps some of the most dramatic displays of variation can occur between the sexes, where males and females can differ so greatly in appearance that they resemble different species. Research in many systems has demonstrated that much of this variation is driven by sexual selection, the force responsible for the evolution of traits that are important for acquiring mates. Individuals may invest as much energy as possible into such sexually selected traits because doing so will give them a competitive advantage for mate acquisition. These traits are therefore considered condition dependent, as their expression is dependent upon the energetic condition of the individual that possesses them. While condition dependence has been the subject of many studies, it is not well known how it may vary between closely related species that share the same traits. If closely related species vary in condition dependence of their shared traits, then this implies that condition dependence could be important for the evolutionary diversity of sexually selected traits.

The rainforest at the La Selva Biological Station in Costa Rica

The lowland rainforest at the La Selva Biological Station in Costa Rica

Together with students from Grinnell College and Reed College, and as part of an OTS (Organization for Tropical Studies) course that I took as an undergraduate at the University of Virginia, we took to the lowland jungles of Costa Rica to answer this question. We studied two anole species from Costa Rica, the slender anole (Anolis limifrons) and the ground anole (Anolis humilis). Specifically, we tested whether several traits that they had in common exhibited condition dependence, including dewlap size, aspects of jaw morphology, and sprinting speed. To test for condition dependence, we first calculated two conventional indices of body condition, the residual index and the scaled mass index, which both take into account an organism’s mass, given its length. We then obtained residuals from the relationship between our variables of interest (dewlap size, jaw width, jaw length, and sprint speed) and snout-vent length (a measure of body length), which allowed us to control for the fact that trait sizes often scale with the overall size of an animal. Finally, we used bivariate linear regressions to test the effect of our indices of body condition on our residual traits of interest, with a significant positive relationship suggesting condition dependence. We found that dewlap size (a trait important for sexual signaling) and jaw width (a trait important for bite force and male combat) exhibited condition dependence in ground anoles, but not in slender anoles. In contrast, neither sprint speed nor jaw length were condition-dependent in either species. Importantly, the presence of condition dependence in one species, but not the other, implies that the condition dependence of shared traits is evolutionarily labile. Additionally, by detecting condition dependence in the dewlap of ground anoles, which have a larger dewlap given their body length when compared to slender anoles, our findings may indicate that the strength of sexual selection differs between these two species. Lastly, our research suggests that variation in condition dependence of the dewlap among species could contribute to the extraordinary diversity of dewlaps in the Anolis genus.

If you would like to read the full paper, published in the Journal of Experimental Zoology Part A, go to:

Signals and Speciation: Do Dewlap Color Differences Predict Genetic Differences?

Dewlap and genetic differences between co-occurring Anolis distichus and A. brevirostris

Dewlap and genetic differences between Anolis distichus and A. brevirostris at sites where they co-occur on Hispaniola.

Here at Anole Annals, we’re all familiar with the replicated evolution of different anole ecomorph types in the Greater Antilles. However, divergence into these different ecomorph classes is not enough to explain how the group became so speciose on these islands. Additional factors must therefore have promoted speciation throughout the history of the group.

One potential factor is the flashy anole dewlap. Dewlap diversification across anoles has led to the remarkable array of dewlap color, pattern and size we see today. If dewlap differences did indeed drive speciation in anoles, or are involved with the maintenance of species boundaries, we might expect that as differences in dewlap color and pattern increases between species, genetic differentiation will also increase through fewer hybridization events.

In our study that just came out in the Journal of Herpetology, Rich Glor, Anthony Geneva, Sabina Noll and I set out to test this using two widespread species from the Anolis distichus species complex, A. distichus and A. brevirostris. These two species co-occur in many locations on Hispaniola and, while they often differ in dewlap color where they do co-occur (yellow with an orange patch vs. all pale yellow), in other areas, they co-occur with similarly pale dewlaps. Using mitochondrial DNA, microsatellite and AFLP data, we investigated patterns of genetic differentiation at four sites: two where the species differ in dewlap color, one where the species share the same dewlap color, and another where pale dewlapped A. brevirostris co-occurs with two A. distichus subspecies (one with a similarly pale dewlap and the other with an orange dewlap).

In general, we found that A. distichus and A. brevirostris looked like “good species,” with strong genetic differentiation and little evidence of hybridization, even at a site where they share the same dewlap color. This suggests that dewlap color differences are not associated with genetic differentiation in a manner one might expect if dewlaps were involved in the speciation process or in maintaining species boundaries. However, at the site where A. brevirostris co-occurs with two A. distichus subspecies with both similar and dissimilar dewlap colors, we found some evidence of hybridization and the species were not as highly genetically differentiated. This discrepancy suggests that site-specific factors could be influencing the dewlap’s role in speciation or maintaining species boundaries. For example, as Leo Fleishman’s and Manuel Leal’s work has shown (e.g. 1, 23), the dewlap’s effectiveness as a signal is dependent on the light environment. Further understanding about the environmental differences among our study sites, how species utilize the available light microhabitats within each site, and how the dewlap looks to anoles at each site could provide more insight into our findings.

On the other hand, perhaps we need to be looking beyond the dewlap and focusing instead on whole signaling displays. Anole behavioral displays can also be strikingly different among species (e.g. 1) and may instead be the key to understanding species diversification in Greater Antillean anoles.

Happening Now: The Latin American Congress of Herpetology

The 11th Latin American Congress of Herpetology is underway right now at the Museo de Zoología QCAZ at Pontificia Universidad Católica del Ecuador. Although I could not attend, I have been following the meeting vicariously as attendees have been using the Twitter hashtag #latinherps to document the meeting.  From those tweets alone, it appears the meeting has featured a series of fantastic talks, including many on anoles. If you are not a Twitter user you can still follow along by clicking more below to see all tweets from the Congress. Finally to Congress attendees, if any of you are interested, it would be great to have you contribute Anole Annals posts (or even comments below) on talks from the meeting.

Continue reading Happening Now: The Latin American Congress of Herpetology

JMIH 2017: Nobody Gets Tired of Looking for Anoles!

Amy Yackely Adams presents at JMIH 2017.

Amy Yackel Adams presents at JMIH 2017.

All anole field biologists have been there, right? It’s the middle of the night, and you’re walking around the forest searching for sleeping lizards in the trees. You’re probably wearing a headlamp, so the bugs are flying around your face, and your eyes start to strain as you get sleepy and you’re entering hour three or four of the search. This searcher fatigue could lead to the kinds of unintentional bias that can interfere with our research. But there’s good news when it comes to anoles, as Amy Yackel Adams, a statistician with the USGS in Fort Collins, Colorado, reported on the last day of JMIH.

Dr. Yackel Adams works with a Rapid Response Team whose goal is to prevent the spread of the worst invasive species. When a report came in of a sighting of a brown tree snake on the island of Saipan (in the Northern Mariana Islands, western Pacific Ocean), the team of experienced herpers deployed to Saipan and began intensive nightly surveys to assess the possibility of a brown tree snake population there.  Luckily, they didn’t find any of these snakes in the surveys, but they did log 20,000+ sightings of other vertebrates! These included emerald tree skinks, several species of geckos, a variety of small mammals, and the green anole (Anolis carolinensis). Dr. Yackel Adams saw an opportunity to use this rich dataset to statistically test for two types of bias that could occur in such surveys – searcher fatigue (both across the 4-hour nightly searches, and across the up-to-31 day deployment), and searcher bias in taxon detection.

The team of 29 searchers covered a total of 387 km of transects during the 31 days, and found a total of 5,800 sleeping green anoles during this time. (Wow!!) In terms of short-term searcher fatigue, there was a slight decrease in tree skink and mammal sightings as the night progresses, and gecko sightings were generally stable over the night, but far MORE green anoles were sighted in the later hours of the night. And over the long term, skinks and anoles were MORE likely to be detected the more nights a searcher worked, and there was no evidence of long-term searcher fatigue.  So, that’s why my take-home message was “nobody gets tired of looking for anoles!”

There was, however, significant taxonomic bias among the searchers – for example, the skink-to-anole sighting ratios ranged from 0.86 to 9.5. Dr. Yackel Adams concluded that this type of bias could be a real problem for certain kinds of studies, and we should be aware that differences among sightings by our survey team members could be potentially problematic in statistical analyses.

If West Indian Weevils Colonized the Mainland 19 Million Years Ago, Were Norops Anoles Along for the Ride?

Exophthalmus scalaris. Credit:

Exophthalmus scalaris. Credit:

In their 2008 review  “Are islands the end of the colonisation road?” Bellemain and Ricklefs (2008) concluded that oceanic islands could be important sources of colonisation of mainland continental areas and cited anoles of the Norops clade as a notable success. There are more than 5 times as many Norops clade species in Central and northern South America as in the West Indies; the 23 extant Caribbean species in the clade are distributed in Cuba and Jamaica with one species in Grand Cayman (Nicholson et al, 2005). Data in Nicholson et al (2005) gave support to the reverse colonisation hypothesis, but did not offer specific dating for the colonisation.

New analyses of 65 species in the Exophthalmus weevil genus complex (Zhang et al 2017) have turned up results that are of significance in understanding the biogeographic history of Caribbean anole dispersal and diversification. Like anoles of the Norops clade, the weevils show reverse colonization (island-to-continent), with diversification on the mainland and diversification within the islands. The data also give some support for overwater dispersal as the factor best explaining ancient between-island distribution.

Zhang et al’s best fit biogeographic model gives an estimate of 19Ma for a jump dispersal of Exophthalmus, most likely from Hispaniola,  which went on to diversify into more than 40 species in Central America.   So – did the anoles and the weevils make their journeys to the mainland around the same time and under similar conditions? Can this weevil study and the techniques it uses to arrive at its conclusions inform anole evolution and dispersal?


Bellemain, E and RE Ricklefs (2008) Are islands the end of the colonisation road? Trends Ecol Evol. 2008 Aug; 23(8):461-8. doi: 0.1016/j.tree.2008.05.001. Epub 2008 Jun 26.   (Correction to citation numbering: Trends Ecol Evol. 2008 Oct; 23(10):536-7).

Nicholson, KE, RE Glor, JJ Kolbe, A Larson, S Blair Hedges, JB Losos (2005) Mainland colonization by island lizards.  Journal of Biogeography 32 (6), 929-938.

Zhang, G, U Basharat, N Matzke, NM Franz (2017) Model selection in statistical historical biogeography of Neotropical insects—The Exophthalmus genus complex (Curculionidae: Entiminae). Molecular Phylogenetics and Evolution, 109, 226-239. DOI: 10.1016/j.ympev.2016.12.039.

JMIH 2017: Brown Anoles Thrive under Artificial Night Light

Chris Thawley presents at JMIH 2017.

Chris Thawley presents at JMIH 2017.

For most of the history of life on earth, the only sources of light at night were the moon and stars. Yet with the invention and rapid spread of electric light, species around the world now face a novel evolutionary pressure: artificial light at night, or ALAN. Artificial light likely has an especially strong effect on animals in city habitats, such as the urban-adapted brown anole lizard, Anolis sagrei. Chris Thawley and Jason Kolbe at the University of Rhode Island set out to determine whether brown anoles were negatively impacted by ALAN.

In addition to their abundance in urban environments, brown anoles are a particularly good species for this study.  Previous studies of brown anoles have shown that photoperiod influences the onset of reproduction at the beginning of the breeding season, and that several behavioral traits change under artificial light. In addition, work by Moore and Menaker has shown that pineal production of melatonin in this species is significantly altered by photoperiod.  So, would ALAN influence brown anole growth and reproduction?

Chris and Jason collected lizards from south Florida and set up a lab experiment where some lizards experienced a normal photoperiod, and others were exposed to ALAN that mimicked landscape lighting. Their results were quite unexpected! ALAN actually increased female growth, resulted in eggs laid earlier in the season, and increased the reproductive output of small females – but did not affect the number or mass of eggs or hatchlings.  And, ALAN females did not exhibit more stress (measured via circulating corticosterone) than control females.

So are brown anoles just superlizards? Do they have no trade-offs that result from ALAN? Chris suggested that it’s possible that such trade-offs may appear in studies over a longer time period, or in traits not measured here. Or, perhaps ad libitum food and the absence of predators in the lab remove the costs of ALAN. Or, maybe these really are indefatigable lizards!

JMIH 2017: Removal of Curly-tailed Lizards Increases Survival of Urban Brown Anoles


Interspecific Interactions Between Two Species of Invasive Lizards in an Urban Environment; Camila Rodriguez-Barbosa and Steve Johnson

An extensive body of work has addressed the eco-evolutionary impacts of the Northern Curly-tailed Lizard (Leiocephalus carinatus) on Brown Anoles (Anolis sagrei) (much of it receiving coverage right here, here, and here on Anole Annals!). These species co-occur not only on many Caribbean islands where much of this research has taken place, but also within the urban matrix of southern Florida, where both species are introduced.

Camila Rodriguez-Barbosa and Steve Johnson investigated the impacts of curlies on brown anoles in shopping centers in southern Florida where both species were plentiful. Camila first collected baseline data on anole and curly populations at eight sites before embarking on a quest to eliminate curlies from four of her sites. Over the next four months, she removed over 300 (!) curlies from these sites, many of which had brown anole remains in their stomachs.

She found that this removal had serious consequences for brown anoles. Compared to anoles from shopping centers where curlies were unchanged, A. sagrei at removal sites experienced higher survival and consequently greater abundances. These anoles also shifted to lower perches once curlies were removed, mirroring results from previous work which show that the introduction of curlies leads to brown anoles occupying higher perches to escape this dangerous predator. Camila’s work suggests that brown anole/curly-tailed lizard interactions may be similar even in very different habitats and provides a fascinating look at lizard life (and death) in the urban sprawl of southern Florida.

JMIH 2017: Brown Anole Reproductive Output Varies Seasonally


Tim Mitchell, Josh Hall, and Daniel Warner: Seasonal Shifts in Anolis sagrei Reproduction Invoke Challenges for Scientific Reproducibility

Sometimes a scientist just needs hundreds of hatchling anoles for an experiment. Tim Mitchell found himself in this position recently, and, like a good lizard ecologist, he started breeding colonies of anoles in the lab to produce eggs to incubate until hatching. As he created three different breeding colonies from brown anoles (Anolis sagrei) in central Florida, one each in February, June, and September, Tim found that he had also created an ideal situation in which to examine how the reproductive condition and output of brown anoles varies across the breeding season.

Tim, along with his coauthors Josh Hall and Dan Warner, found that females produced eggs with significantly greater mass later in the breeding season. These eggs took longer to produce than those earlier in the year (a greater interclutch interval), and the eggs resulted in hatchlings that had higher mass in relation to the weight of their eggs. These reproductive differences remained even after accounting for the fact that female anoles were also larger and heavier later in the year.

These findings suggest that female A. sagrei may shift their reproductive effort from producing a higher quantity of eggs (i.e., more, smaller eggs resulting in smaller hatchlings) in the beginning of the breeding season, to producing higher quality eggs (i.e., fewer, larger eggs resulting in larger offspring) later in the breeding season. Tim’s findings also stress the importance of investigating and accounting for seasonal differences when examining reproductive output in lizards.


JMIH 2017: Costa Rican Anole Ecology


Greetings from Austin, Texas, and the Joint Meeting of Ichthylogists and Herpetologists! Chris Thawley and I have appointed ourselves to be your AA reporting team from JMIH, and we’re aiming to post updates from each of the 8 anole talks and posters at this meeting.

Brian Holt

Brian Folt

On the first day of the conference, there were two exciting talks on the ecology of Costa Rican anoles. The first was by Brian Folt, a graduate student in Craig Guyer’s lab at Auburn, who developed a model of predator-prey co-occurrence where one of the prey were anoles (Anolis (Norops) humilis) and the predators were…spiders?!  Yes, wandering spiders, or ctenids, can prey upon the small anoles on the forest floor. (The other putative prey were poison dart frogs, who have a relatively similar life history to anoles.) Brian performed an extensive field study in 14 plots at La Selva Biological Station, conducting visual encounter surveys for anoles, frogs, and spiders, and recording arthropods in leaf litter samples. He used two-species occupancy models to determine how prey were affected by the presence of the predator and by resource abundance in the leaf litter. The result? Anole occupancy was lower where spiders were absent, and the detection probability of anoles was higher when spiders were present and detected. This suggests that anoles are responding behaviorally, such that they may increase their vigilance when predators are around.

Michelle Thompson

Michelle Thompson

I then ran across the conference center to catch the next anole talk – a terrific presentation by Michelle Thompson, a graduate student in Maureen Donnelly’s lab at Florida International University. Michelle studied whether thermal quality differed across the stages of forest succession, and how that affected Anolis (Norops) humilis and A. (N.) limifrons distributions. She worked across transects of pasture, secondary forest, and old growth forest in both upland and riparian sites. Michelle measured the thermal quality of each habitat, the thermal preferences of the lizards, and the location and abundance of the lizards. She found that thermal quality was lowest in the pasture sites, as temperatures were frequently higher than the lizards prefer. Yet, in these pasture sites, riparian habitat with remnant trees provided a thermal refuge for the lizards. This kind of work can help us understand why and how species may respond differently to human-caused alterations in habitat structure and temperature in our changing world.

Stay tuned for updates from JMIH, and follow the #JMIH17 hashtag on twitter for more herp-related news!

Evolution 2017: Anoles and Ameivas Have Similar Gut Microbiomes

Late Breaking: one last Evolution 2017 post!  Last weekend during the Evolution meeting, I had a chance to chat with Iris Holmes (Ph.D. student, University of Michigan) about the poster she presented. Initially not on our watch list because of the lack of “anole” in the description, my eye caught the dewlapping lizard perched at the top of her poster from across the room.

2017-06-25 19.53.04

Iris presented her work on gut microbiomes of two groups of lizards: anoles and ameivas. She wanted to know if different taxa have different gut microbiomes and to what extent diet influences bacterial composition of gut microbiomes. Her collaborator (Ivan Monagan) collected scat samples from 22 Anolis dollfusianus and 9 Ameiva from an agricultural area in the Soconosco region of Chiapas, Mexico. Together, they then sequenced both the gut bacteria and the digesting prey with two 16S primers. Iris chose to target the prey as well because she wanted to know if they were eating different things and how different stages of digestion influence gut bacteria communities.

Iris found that there were no clear differences between the gut microbiomes of anoles and ameivas. Both species had gut microbiomes dominated by three main phyla: Proteobacteria, Firmicutes, and Bacteroidetes. Little is currently known about how these bacteria relate to digestion and health in reptiles, but Iris commented that we can make some guesses based on studies in other taxa. Proteobacteria are a disease indicator in mammals, but appear to be normal in reptiles and birds. Firmicutes and Bacteroidetes are both important for digestion of carbohydrates and fats (respectively) in mammals. Iris found that there was a loose correlation between the amount of prey consumed and the abundance of Bacteroidetes, suggesting these bacteria also play a role in digestion in lizards. She also found that there was an apparent tradeoff between the Proteobacteria and the two other groups – sequence abundance of proteobacteria was negatively correlated with abundance of Bacteroidetes and Firmicutes. Overall, this is an interesting first step in understanding the gut microbiomes of reptiles and how they differ (or don’t) between groups.

Metabolism Rate Data on Anoles?

I’m hoping that some of you out there have been collecting Basal Metabolic Rate or Resting Metabolic Rate data on Caribbean anoles!

I’m working with a group of scientists on a large-scale comparative database on circulating hormones in free-living vertebrates – we call our collaboration HormoneBase – and we’re hoping to look at relationships between hormone levels and metabolism. (We’ll be presenting some of this work at the Society of Integrative and Comparative Biology meeting in January 2018 – check out our symposium announcement here!) We have a good list of anole species in the database, thanks to the work of Jerry Husak and Matt Lovern (2014), but it seems that very little metabolism rate data are available for these species. Do you know of such data, or do you have them – published or unpublished? If so, please contact me (!



Husak JF and MB Lovern. 2014. Variation in steroid hormone levels among Caribbean Anolis lizards: endocrine system convergence? Hormones and Behavior 65:408-415.