Anoles of South America Part 1: Diversity North of the Andes

Although anoles are much more well studied and understood in the Caribbean and United States, there is a decent amount of diversity of these lizards throughout South America as well. In consultation with Jonathan Losos, I have agreed to write a series of posts outlining the South American diversity (parts 1 and 2), invasives (part 3), and conservation (part 4) of these animals. This post (diversity part 1) will describe the species that occur in the regions of South America north and west of the Andes along the Caribbean and Pacific Coasts, respectively, while diversity part 2 will focus on the Amazon and everything south and east of the Amazon.

In case you don’t know me, I had experience as a graduate student studying lizard functional morphology of toes and tails in North, Central, and South America. More recently, I have become way too involved in identifying lizards on iNaturalist.org, with a focus on lizards of the New World. To satisfy my own particular obsessiveness, I have tried to identify every lizard observed in the Amazon (all 300+ species of them!) and have expanded those lizard-identifying efforts into other areas as well. It was these efforts that gave me the idea that perhaps the AnoleAnnals might be interested in what I’ve noticed from this relatively understudied part of the world. As such, I will draw liberally from data available on iNaturalist to illustrate my points.

Anole diversity along Caribbean and Pacific Coasts

As of this writing, there are currently nearly 12,000 observations on iNaturalist of 102 species Anolis in South America (“research grade” observations filtered by “wild” and “has photos”). Compare this to the 238,000 observations of 270 species from North America (on iNat this region includes all of Central America, Mexico, the United States, and the Caribbean). Only eight of the species of South American anoles are introduced non-native species (more on this in part 3). This suggests that there are nearly 100 species of Anolis endemic to South America. Clearly, the anoles from South America are not as diverse and not as well observed as those from other areas.

Anoles along the Pacific Coast range from southern Ecuador (and perhaps northern Peru, but not well documented) north to the junction with Central America and continue along the Caribbean Coast east to the mouth of the Orinoco River in Venezuela (note: according to iNaturalist, Guyana, Suriname, and French Guiana are part of the Amazon region even though they may not be part of the Amazon watershed). This region is separated from the Amazon by the Andes Mountains and Orinoco River and very few species cross these particular barriers. Hence, many of the anoles of northern South America still have some degree of endemism and it is this endemism (and the resulting beta diversity) that I highlight below. Keep in mind that new species are being described on a regular basis, so the list below is almost certainly incomplete, but still illustrates my general point.

The species of Anolis that occur in north and west of the Andes include (organized by their geographic distribution):

Western (non-Amazonian) Ecuador (n=9)

A. bionotatus, A. bitectus, A. fasciatus, A. festae, A. lyra, A. neomonteae, A. parilis, A. peraccae, A. proboscis

Western (non-Amazonian) Colombia (n=18)

A. anchicayae, A. antioquiae, A. antonii, A. calimae, A. danieli, A. eulaemus, A. gorgonae, A. heterodermus, A. maculigula, A. macrolepis, A. mariarum, A. medemi, A. quimbaya, A. solitarius, A. sulcifrons, A. tolimensis, A. urraoi, A. ventrimaculatus

Western Ecuador/Colombia (n=13)

A. aequatorialis, A. anchicayae, A. chloris, A. dracula, A. fraseri, A. gemmosus, A. gracilipes, A. granuliceps, A. limon, A. lynchi, A. maculiventris, A. parvauritus, A. princeps

Northern Colombia/Venezuela (n=3)

A. jacare, A. onca, A. tetarii

Northern Venezuela (n=4)

A. anatoloros, A. annectans, A. squamulatus, A. tigrinus

Aruba, Bonaire, and/or Curaçao (A-B-C islands) (n=2)

A. bonairensis, A. lineatus

Found in western Colombia as well as crossing the Darién Gap into Central America (n=8)

A. biporcatus, A. gaigei, A. latifrons, A. poecilopus, A. purpurescens, A. triumphalis, A. tropidogaster, A. vittigerus

Along Caribbean Coast as well as crossing into the Amazon (n=2)

A. auratus, A. planiceps

Non-native anoles (n=8)

A. aeneus, A. cristatellus, A. marmoratus, A. porcatus, A. richardii, A. sagrei, A. trinitatis, A. wattsii

Several things become apparent when considering the above diversity.

—First, there are roughly 67 species of anoles that occur north of the Amazon. Given that there are only about 102 anole species that occur in all of South America, perhaps 2/3 occur in the area adjacent to the speciose Caribbean. That leaves only 1/3 of the South American anole species occurring south and east of the Andes Mountains.

—Second, of the 67 anole species found north of the Amazon, 47 species (70%) are endemic to this region of mainland South America, two species (3%) are endemic to the A-B-C islands, eight species (12%) cross from mainland South America into Central America, and two species (3%) are found extensively in the Amazon as well. So, not as diverse as North America, but with outstanding regional endemism. This regional endemism translates into an impressive beta diversity with geographic replacement probably the rule rather than the exception. In other words, when travelling from one valley to the next, species replacement (beta diversity) occurs at a relatively high rate so that nearly an entirely different set of species may be present in adjacent areas, depending on the nature of the barrier between them. Making this even more of a challenge, in some instances the barrier appears to be the low-lying valley instead of a mountain ridge. For example, in Colombia, Anolis heterodermus Duméril 1851 was recently split into eight distinct species (Morena-Arias et al. 2023). Both the various mountain ranges (Cordillera) as well as the Magdalena River valley appear to act as barriers isolating the various lineages into distinct species. Hence, in terms of trying to identify the Anolis in some of these areas, accurate geographic location is a must.

—Third, there are only eight species (12%) of non-native anoles in all of South America (including the A-B-C islands and Trinidad and Tobago). All eight occur in northern South America and only one or two (A. porcatus and perhaps A. sagrei) occur elsewhere in South America. Contrast this with the 16 or so species of introduced anoles in North America, which have made extensive inroads in places like the southeastern United States, southern Mexico, and certain of the Caribbean Islands. These eight introduced species will be the focus on part 3 of this series.

One of the interesting things about the high degree of endemism of the anoles of northern South America is how quickly it probably occurred. Andean orogeny began perhaps 20 million years ago (mya) and lasted until about 8 mya. Hence, the diversity differences between the occidental (western) and oriental (eastern) regions of countries such as Ecuador, Colombia, and Venezuela are notable for their near completeness. For comparison, the Laramide orogeny that created the Rocky Mountains occurred between 80-55 mya; the Alleghanian orogeny that created the Appalachian Mountains occurred between 325-260 mya. Hence, the temporal arrival of the Andes created spatial barriers to gene flow and geographic complexity that is reflected in the high beta diversity of Anolis both within and between regions of South America.

In conclusion, northern South America is the diversity hotspot for anoles. The anole species north of the Amazon are dominated by native, highly endemic species that often have geographic ranges limited to a single country, a single valley, or even one slope of a single mountain ridge. Moreover, there are relatively few introduced anoles in South America with most of those introduced to the A-B-C islands and Trinidad and Tobago. Given that almost no species of anoles cross the Orinoco River or Andes Mountains and very few species occur extensively in the Carribean or Central America, the anoles of northern South America represent a fairly separate and distinct fauna when compared to other geographic regions within the distribution of Anolis.

 

Literature cited:

Moreno-Arias, R. A., Méndez-Galeano, M. A., Beltrán, I., & Vargas-Ramírez, M. 2023. Revealing anole diversity in the highlands of the Northern Andes: New and resurrected species of the Anolis heterodermus species group. Vertebrate Zoology, 73, 161-188

Profile Photos of Green Anoles Needed

Hello fellow anole enthusiasts!

I’m reaching out because I’m in need of some specific anole photos for my research. I’m currently wrapping up my dissertation on the diversification of the Anolis carolinensis subgroup, and I would like to include head profile photos of male anoles in their light phase with the dewlap fully extended (see figure).

I’m specifically looking for images of the following species:

Anolis brunneus

Anolis carolinensis

Anolis longiceps

Anolis maynardi

Anolis smaragdinus

If you have high-quality photos of any of these species, I would greatly appreciate your contribution. All photos will be fully credited to their respective authors, and I’d be happy to share a PDF copy of the published article once it’s completed. Please contact me at javiertorres@ku.edu.

Thank you in advance for your support!

Best regards,

Javier

Through Local Eyes: Unveiling the Biology of Anolis biporcatus with Citizen Science

Photo credit to Francisco Araya.

Written by Wagner Chaves-Acuña and Diego Salas-Solano

Recent fieldwork at the stunning Veragua Rainforest Reserve in Costa Rica’s Caribbean region has brought to light fascinating new insights about Anolis biporcatus, a vibrant green lizard that spends much of its life hidden high in the forest canopy, far from human observation. Through the combined efforts of dedicated scientists and the invaluable local guides and staff at Veragua, we’ve managed to document behaviors never before recorded in this species (see Chaves-Acuña & Salas-Solano, 2024).

Collaboration is Key.

This work wasn’t accomplished by scientists alone. Our collaboration with the naturalist guides and community members at Veragua has been a key part of the process. What might seem like a groundbreaking revelation for us is often part of the daily lives of these dedicated people who live alongside these and many other forest species. Guides such as Kenneth Gonzales Najera, Francisco Araya, Mónica Rojas Rodríguez, and Junior Calderón Villalobos have contributed enormously by observing and capturing footage of the lizards’ behavior, helping us document crucial moments. Thanks to the proximity of the research station to the field sites, we are able to quickly validate and build on this information, creating a dynamic exchange of knowledge, where both science and local expertise come together to unveil nature’s mysteries.

This connection with the local community not only enhances our research but also turns them into passionate guardians of the rainforest. By participating in these projects, they become advocates for conservation and sustainable practices, helping protect the unique biodiversity of this incredible region.

Observing a Miniature World Where Every Detail Counts in the Delicate Dance of Attraction.

Visual displays are a fascinating form of lizard communication, especially during courtship and mating. Males, with their striking blue-centered, red-bordered dewlaps and females with their uniformly white dewlaps both engaged in frequent dewlap extensions during courtship. When a male flares his dewlap, he’s sending important signals about his strength, health, and even genetic fitness, helping females make informed choices about their potential mates. On the other hand, the female’s dewlap display may signal her receptivity to courtship or convey her status to other lizards of the same species. In some cases, it may even function as a way to deter unwanted advances or to assert her presence in a particular area.

The Magic of Color Change.

These lizards normally remain bright green to blend in with the leaves and branches around them. But during courtship, the males undergo rapid color transformations, shifting to darker tones with bold stripes or patches. This color dance may be a visual signal to the females or perhaps a warning to other males in the area. In one particularly dramatic moment, we witnessed a male change color just after biting the female’s neck, a gesture that often precedes mating.

Mating isn’t Easy When You Live in a Forest of Towering Trees.

Most of the time, we observed the lizards mating head-down on vertical tree trunks. The male would hold on tightly to the female with one arm and leg, wrapping the others around her midsection to ensure they stayed in contact. Sometimes, the male’s tail even helped him maintain balance. It’s amazing to see how well they’ve adapted to life in the forest’s complex, vertical world.

What’s particularly surprising is that, although Anolis biporcatus is usually found high up in the canopy, most of our observations happened at just 2 meters above the ground, with only one at 3 meters. This suggests these lizards are more flexible than we thought, using a range of heights in the forest to suit their needs.

Diet Notes on Beautiful Blue Knight Anole

Photographs by Odey Martínez Llanes

Read all about it!

Photograph by Odey Martínez Llanes

Anoles Provide Ecosystem Services

Photo by Julie Katz

Hard to imagine a more important service than this! Miami resident Julie Katz provides the details: “When I walked out of my condo building one night this past spring to walk my dog I encountered a first – a small lizard feasting on a roach.”

Go anoles!

Non-native Herpetofauna of Aruba: the Art and Science of Documenting Species Invasions

Anolis porcatus (left) found at a plant nursery in Aruba (right).

It was late December 2018. My partner in work and life, Matt, and I were headed back to Curaçao to conduct some follow-up sampling. We had recently completed a field season in Curaçao where we had discovered three newly introduced gecko species (Behm et al. 2019).  Since we had to fly through Aruba to get to Curaçao, we decided when booking our trip to spend some time in Aruba, too, to look around for introduced herps, since that’s one of our favorite ways to spend our time when traveling (we’ve even been known to trick family members into searching for introduced herps while on vacation – e.g., Behm et al. 2018). Aruba and Curaçao are neighbors and since Aruba is even more economically connected than Curaçao, we suspected there may have been undiscovered non-native herps there.

What we did not expect was that in less than 3 days on Aruba, we would find three new non-native species (Anolis gingivinus, A. cristatellus and Hemidactylus frenatus) and document a range expansion for a 4th non-native anole (A. porcatus) already known to be on Aruba (Behm et al. 2022). Even though we focused our surveys on properties like plant nurseries and resorts that had a strong likelihood of harboring introduced species due to their influx of ornamental plant shipments, this was still a very high number of species in a short amount of time.

I wanted to understand how these newly documented species fit within the context of the other non-native herps on Aruba. Specifically, I wanted to understand the circumstances surrounding the initial introductions and the current habitat use of the species on Aruba to assess whether any emergent patterns could be gleaned for Aruba. If so, these patterns may lend further insight into how and why the Caribbean has emerged as a hotspot for species invasions (e.g., Helmus et al. 2014, Perella and Behm 2020, Gleditsch et al. 2023).

What seemed like a simple task ended up being more challenging than expected due to the scant and sometimes contradictory information in the literature surrounding the species introductions. To make sense of it all, I enlisted the help of a brilliant undergraduate in my lab, Gianna Busala, whose tenacity and attention-to-detail made it possible to reconstruct some of the invasion history on Aruba.  Our recent publication (Busala et al. 2024) compiles all of what we found about the introduced herps on Aruba.

In conducting our research, there were several issues we encountered.

Mississippi Kite Eats Green Anole

Photo by Christa Denning (aka, @DenningDesign)

From Twitter

Lanceolated Monklet Eats a Slender Anole

Read all about it in Herpetology Notes.

How Do Body Color Changes Affect an Anole’s Conspicuousness to Conspecifics and Predators?

Water anole from the Osa Peninsula, Costa Rica (Photo by Lindsey Swierk)

Rojo A, & L Swierk. 2024. How does rapid body color change affect the conspicuity of lizards to their predators and conspecifics? Behavioral Ecology and Sociobiology 78: 78.

 

Written by Andrés Rojo:

The water anole (Anolis aquaticus) is a Costa Rican anole that changes color and pattern in different microhabitats as a form of camouflage to avoid detection by their predators, which include birds like motmots and kingfishers. Because of its body color complexity and rapid color changes, water anoles are great study organisms to use when examining how animal camouflage, social signaling, and visual perception interact.

My research project was inspired by Dr. Lindsey Swierk and her lab’s work on water anole color change and camouflage. I joined the Swierk lab as an undergraduate in February of 2021, as I was interested in tropical ecology and animal behavior. I am also a photo hobbyist and experiment with color and full spectrum photography. Dr. Swierk thought that I would be a good fit with the research team studying water anole colors and patterns. When she told me about it, I was motivated by my enthusiasm for photography and wildlife research to take on the project, especially the idea that I could modify research-grade photos to model how animals see one another.

An example color cloud map showing the colors of a water anole (gray) and its substrate (red) as seen through an avian visual system. The X axis represents a green-to-red progression of colors, and the Y axis represents yellow-to-blue colors. Water anoles do not have UV body colors in the regions measured (dorsal and lateral surfaces). Darker tones indicate more pixels of colors at that location within chromaticity space.

Dr. Swierk and I decided to test whether the body coloration of A. aquaticus would be perceived differently by the visual systems of water anoles’ predators and their conspecifics. We used ImageJ and the micaToolbox (QCPA) to model the two visual systems and apply them to each photo taken in the field. The anole and substrate photos were converted into color maps, which could be compared to determine how much the color of the anole overlapped with the color of the substrate in the visual systems of predators and conspecifics – a measure of conspicuousness of anoles according to both visual systems.

A comparison of body color – background color overlap of water anoles as seen through bird and anole visual models, including males (blue) and females (gold).

Although we found no significant difference in how camouflaged A. aquaticus were perceived by the visual systems of predators and conspecifics, our results suggested that males more consistently color matched their microhabitats compared to females and that females were less likely to color match their backgrounds in their lighter phase, suggesting a sex difference in preferred conspicuity in more exposed habitats. These could present topics for future research into how anoles use color camouflage to avoid being detected by predators.

Limited Morphological Differences of Brown Anoles (Anolis sagrei) between Their Native Cuban and Invasive Florida Range

Jars containing museum specimens are on a cart next to the shelf.

The ubiquity of Brown Anoles (Anolis sagrei) in Florida means that they are constantly on my mind. Given the proposition that rapid evolution may be important to invasive species success, I was curious to determine if invasive Brown Anoles in the southeastern United States, centered in Florida, have any morphological differences from Brown Anoles in their native range in Cuba. To address this question, I measured museum specimens to compare Brown Anole morphology between their invasive and native range and “go back in time” to see if Brown Anole morphology has changed since their invasion.

As reported in our recent paper, Brown Anoles have broadly similar morphology between their native Cuban and invasive Florida ranges. Additionally, we found no clear evidence of the measured morphological traits changing over time. These results suggest that rapid morphological evolution may not be essential to the success of invasive Brown Anoles.

Abstract:

Understanding why some species and not others are successful global invaders is an important question in ecology and evolutionary biology. There is much debate on the role that rapid post-invasion adaptation plays in the success of invasive species. Here, we investigated signals of rapid and broad-scale morphological evolution in Anolis sagrei (Brown Anole) between their invasive and native distributions. Although we found significant differences in a few morphological characters between invasive and native Brown Anoles, the morphological variation present in the species broadly overlapped between both populations and has not significantly changed over the last century. These results suggest the invasive success of Brown Anoles in Florida may not be due to major evolutionary change from their Cuban ancestors.

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