Australian Cryptoblepharus: Convergence across a Continent

When it comes to adaptive radiations, the diversification of Anolis is one of the most striking examples. While Anolis is therefore a well-known rock stars in the league of adaptive radiations, in this blog post I would like to introduce you to one of the mere mortal examples; skinks from the genus Cryptoblepharus.

Cryptoblepharus skinks (“Cryptos”) are small diurnal lizards that have rapidly diversified and are known for their widespread distribution with species present in the Malagasy region, on the Australian continent and on many island archipelagoes in the Indo- and wider Pacific. Furthermore, species that occur on similar substrates are notoriously difficult to identify based on morphological characteristics and a more accurate estimate of species diversity has only recently been accomplished using a widespread genetic screen with allozyme markers (Horner & Adams, 2007).

Distribution of Australian Cryptoblepharus and the three habitat specialists. (a) Topographic map of Australia with the mean point of each species’ distribution plotted and coloured according to habitat type (for complete distribution maps, see Horner & Adams (2007). In situ photographs of (b) arboreal, (c) littoral, and (d) rock specialists (green, blue, and red dots on the topographic map, respectively).

Distribution of Australian Cryptoblepharus and the three habitat specialists. (a) Topographic map of Australia with the mean point of each species’
distribution plotted and coloured according to habitat type (for complete distribution maps, see Horner & Adams (2007)). In situ photographs of (b) arboreal, (c) littoral, and
(d) rock specialists (green, blue, and red dots on the topographic map, respectively).

Whereas species within the same habitat are highly cryptic, species that occur on different substrates (‘rock’, ‘trees’ or ‘beaches’) are relatively easy to distinguish. Rock Cryptos for example, traverse the red sandstone escarpments that are iconic for the Australian outback (think ‘Uluru like’ in terms of color and rock type) and they look very different from Cryptos that occur in a more mesic or coastal habitat. In a recent paper (Blom et al., 2016) we focused on the Australian radiation and explored whether habitat specialization explains current patterns of phenotypic variation in ecologically relevant traits. Using a comparative approach, we quantified the presence of distinct adaptive peaks, the frequency of shifts between such peaks and ultimately discuss the role of ecology in promoting continental radiation.

Anolis Photos from Cuba: ID Help Needed

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I spent a few weeks on Cuba in February-March, and photographed a bunch of different anoles, but I have no way of identifying them. I put all photos on one page with very tentative captions. I’ll appreciate any comments/corrections.

Here are some of the uncertain ones:

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Brown Anoles Invade New Orleans: What Will Happen to the Greens?

Just in time for the American Society of Ichthyologists and Herpetologists meeting in New Orleans next week. From the New Orleans Advocate:

It flashed across the walkway like a lightning bolt, so fast that Bob Thomas had to do a double take. In that split second six months ago, he knew they had finally arrived.

“I’d been waiting for them to arrive in my neighborhood in Metairie. What I saw moved too fast for what we’re used to around here,” said Thomas, a herpetologist who taught at Loyola University and served as the founding director of the Louisiana Nature Center.

“It could only be one thing: a brown anole, Anolis sagrei.”

You’ve seen them — the speckled brown lizards that come out of nowhere and streak across the sidewalks. They travel in hordes — tiny, large and everything in between. Careful! You’re liable to step on them if you don’t pay attention.

Thomas’ neighborhood is far from being the first to experience an invasion of brown lizards. But where did they come from? Why are they so plentiful?

“Brown anoles are an invasive species, not native to the United States,” said David Heckard, curator of reptiles and amphibians at the Audubon Institute. “They are natives to Cuba and the Bahamas and first appeared in the U.S. in Florida. From Florida, they’ve been slowly expanding their range across the Gulf Coast. They’re aggressive and competitive and have even been spotted in Taiwan. They hitch rides on plants and are spread inadvertently by plant nurseries.”

The brown anole looks a lot different than the sleek green lizards we grew up with here in New Orleans (Anolis carolinensis). Generally, A. sagrei has a more compact physique and a shorter skull. A prominent hump appears where muscles attach at the back of the skull. When the brown anole extends its orange and red dewlap (the skin flap below its chin), it looks ferocious, indeed.

By contrast, the green anole looks far friendlier, even when its rosy-hued dewlap is extended. Native to the southeastern parts of the United States (although DNA studies suggest they originated in Cuba and came here a couple of million years ago), green anoles range as far north as North Carolina and as far west as Austin, Texas. They have delicately shaped heads and long, lean bodies. They were once plentiful in New Orleans, but sightings are becoming rare.

So, are the brown anoles killing off the green anoles, fighting over territory and winning? Consuming the green anole’s food supply?

“The theory is that the brown anoles are displacing the green anoles but not necessarily replacing them,” Heckard explained. “It’s believed that green anoles are more arboreal than brown anoles, which are more terrestrial. So, green anoles are being pushed to higher elevations — up into trees and the like. It may seem as though there are fewer of them, but they’re present — you just can’t see them hiding in the leaves and up in trees.”

Simon Lailvaux, a professor in UNO’s department of biological sciences, has studied anoles since working on his doctorate and supports the displacement theory.

“In the Caribbean, where there are dozens of species of lizards, they have learned to partition the habitat and have evolved to live in a specific part of it,” Lailvaux explained. “Green anoles there are trunk/crown inhabitants, whereas brown anoles are trunk/ground inhabitants. Over the millions of years that green anoles have been in the United States, they evolved to be able to occupy the ground because they didn’t have any competition for it. So, the relatively recent invasion of brown anoles has simply forced them back up into trees where they originally lived.”

Are we sure about that? Is anybody counting?

“How can you count green lizards way up on tree trunks and in the leaves at the crowns of trees?” answered Lailvaux. “You can’t.”

According to all three scientists, both types of anoles eat the same things: insects and other invertebrates. There are plenty of those to go around here, so it’s improbable that the green anole’s food supply is in jeopardy. Luckily for the green anoles, they may have a significant competitive advantage over the invaders.

“Brown anoles are cold sensitive and can survive only in a limited temperature range. That means the population of brown anoles crashes when we get a hard freeze, and it takes forever for their numbers to recover,” Lailvaux said. “The green anole, on the other hand, has evolved to be able to withstand lower temperatures, so they won’t be bothered by a freeze. We’re seeing, though, that it is taking less and less time after a freeze for the brown anoles to recover, which means they’re already beginning to adapt.”

The mild winters of the past few years may account for the explosion in the visibility of the brown anoles. But if A. carolinensis is being replaced (not merely vertically displaced) by A. sagrei, it would be a case of a native species dying out because an invasive species outcompetes it. Should we be looking into how to reverse that trend?

“The green anole may be a nostalgic favorite, but we don’t know yet what impact the proliferation of the brown anole will have on it or on other species. The sense is, however, that it won’t be wonderful,” Thomas said.

We know too well what an invasive species can do: Witness the nutria. By consuming the marshes, the animals not only reduced storm surge protection for our area but caused the demise of other species that called the marshes home, Thomas pointed out. Without further study, there’s no way to predict if the success of the brown anole could be similarly dire for the green anole and for biodiversity.

Anole Fabric

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I’m not sure I like anoles being referred to as “ditsy,” but here’s a great opportunity to create lovely anole-wear, not to mention anole curtains, anole quilts and all kinds of other anoliana.

anole fabric

Prince of Thar–Sands of Time…

An adult nipping grass in the morning; notice the bluish tinge on the inner side of the thigh and dorsal part of tail.

An adult nipping grass in the morning; notice the bluish tinge on the inner side of the thigh and dorsal part of tail.

Here is Saara hardwickii , spiny tailed lizards. I observed these lizards in their natural habitat, in the Thar desert in Indian state of Rajasthan. It’s a medium-sized lizard which dwells in semi-arid to arid landscapes of northern India, Pakistan and some regions beyond. A drab colored lizard with a pug head and a distinct fleshy and spiny tail.

Habitat fragmentation and hunting for its tail is the main reason for its dwindling numbers. Folklore has it that its tail has aphrodisiac powers, so its tail is cut and ‘oil’ extracted from it and consumed for the intended purpose.

Interestingly, like iguanas, these lizards also live in a social structure, a ‘society’ composed of adults as well as young ones. They live in ground burrows or termite mounds. Spiny-tailed lizards are diurnal; their activity starts around early morning sun and when the sun sets, surprisingly not even a single individual can be seen! A considerable ontogenic shift in dietary inclination towards herbivory can be seen. Adults feed on grass or diminutive terrestrial flora, whereas young ones are omnivorous, feeding on arthropods.

This fellow was just out of its home and carefully observing its habitat.

This fellow was just out of its home and carefully observing its habitat.

Green Anole Hunting Brown Anole–Foiled

And he’s not happy about it! Photo by Karen Cusick

Details on Daffodil’s Photo Blog.

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Nocturnal Behavior in the Green Anole

I’m currently reading a 274 page tome called “The Biology and Biodemography of Anolis carolinensis” by Robert E. Gordon. Dating back to 1956, this impressive piece of scholarship is Gordon’s Ph.D. thesis. Gordon collected the bulk of his data in biweekly nocturnal surveys of the demography and spatial ecology of two populations of green anoles. The surveys continued for over a year, and consequently, this document is filled with insights into these lizards’ ecology.

One sentence that caught my attention was this, from page 195:

Anolis activity is primarily diurnal, although movement and feeding were observed at night under conditions of bright moonlight.

We’ve had observations of anoles feeding at artificial lights before, but have any of you night-owl herpers observed something similar under natural light?

A figure from Gordon (1956). Can we bring back this elegant asymmetric bar graph plotting style?

A figure from Gordon (1956). Can we please bring back this elegant asymmetric bar graph plotting style?

 

Three-Legged Green Anole

IMG_0873 (2)There are several previous posts concerned with lizards missing feet or limbs (1, 2, 3). At the risk of being monotonous, here is another. I caught this male green anole (Anolis carolinensis) in Auburn, AL this morning (6.24.16). He was sitting on someone’s porch railing at my apartment complex. In addition to his front left limb he was missing 2 fingers on the front right hand and one on the back left foot. He has no other signs of damage, however, not even any evidence of a regenerated tail. I think what sets this example apart from ones previously given is that the entire limb is missing. There is only a tiny nub of bone at the shoulder, and a small flap of skin. Interestingly, the tiny nub moves back and forth beneath the skin when he runs, as if the entire limb were still present and useful.

We (Warner Lab) have had lizards hatch without limbs in the past. We even had one (A. cristatellus) hatch this year with 6 limbs (three front right arms). It is possible that this carolinensis never developed this limb to begin with; however, the tiny flailing nub and flap of skin make me feel that is not the case. Anyhow, this guy seems fat and happy and still moves pretty fast, despite the handicap.

Evolution 2016: Polar Vortex Revisited

Shane Campbell-Staton giving his talk at Evolution 2016

Shane Campbell-Staton giving his talk at Evolution 2016

We’ve heard about the effects of polar vortexes here on Anole Annals before. The infamous 2013/2014 event brought record-breaking snow and low temperatures to the Southern U.S., leaving people and animals both a little chilled. This created the perfect opportunity for Shane Campbell-Staton to investigate the effects of such extreme events on thermal tolerance of the native Carolina Anole, Anolis carolinensis. Shane also spoke about this at SICB earlier this year, and AA contributor Martha Muñoz covered the talk pretty thoroughly here on Anole Annals. Nevertheless, I’ll summarize some key points here in case you missed it.

carolinensis frozen

An unlucky lizard during the polar vortex snow storms in the South.

Shane got lucky in the sense that he had measured thermal tolerance in August 2013 for populations affected by the polar vortex, 5 months before the event. Typically, the cold arctic air is tightly constrained around the North pole, but periodically the boundaries weaken and the cool air expands southward. These events are not regular, so Shane had no idea one was coming that winter or that it would extend so far south. It was serendipitous that his study populations, 3 in Texas and 1 in Oklahoma, were impacted by the extreme weather event. This species, particularly in the Southern portion of its range, is not used to low temperatures and reports came in of anoles dying off during the storm.

Air temperatures for January 5-7, 2014, compared to the 1981-2010 average. Map by NOAA Climate.gov

So Shane returned in August of 2014 and sampled again, curious as to how this cold impacted thermal tolerance. He found that tolerance to low temperatures, measured as critical thermal minimum (CTmin), was lower in some populations after the event! Even more, the difference was greatest in the Southernmost population (Brownsville, Texas). Shane returned again in the fall of 2014 to see if this effect persisted or if it was simply a plastic response to the event. He found that the populations sampled in 2014, and presumably their offspring, still had lower critical thermal minimums. This result suggests that the extreme cold weather had caused an evolutionary shift in cold tolerance via natural selection: only the animals that could tolerate the cold temperatures survived and passed on their cold-tolerance genes. Shane went on to conduct a common garden study to verify that the trait was not simply a plastic response. He found that the lower CTmin persisted in lab-reared animals: strong evidence that these shifts had a genetic basis.

Lastly, Shane looked at the functional genomics of cold tolerance. Using liver tissues to obtain transcriptomes (representing expressed genes), he found several gene modules associated with thermal tolerance including some associated with respiratory electron transport chain, lipid metabolism, carbohydrate metabolism, and angiogenesis/blood coagulation. He also found that the gene expression patterns in the Southern populations affected by the storm resembled the Northern populations that more regularly experience cool temperatures, indicating a common genetically based adaptive response across populations.

Evolution 2016: Using Genomic Tools to Explore Selection and Evolution in Anolis Species

imageBy Pavitra Muralidhar

Adaptive radiation is one of the most intriguing processes in evolutionary biology, and anoles are one of the well-studied examples of this process. Anoles have diversified into over 400 species across the Caribbean and Central America, and contain a multitude of highly divergent morphological and behavioral types. Thanks to an impressive history of research on this clade, we now know quite a lot about the phenotypic aspects of this adaptive radiation; however, we still don’t have a good understanding of the genetic mechanisms underlying this diversity of form, physiology, and behavior. The recent advent of next-generation sequencing, and thus the ability to quickly sequence entire genomes of non-model organisms, offers a tantalizing possibility for investigating the genetic basis of adaptive radiation in Anolis.

Tollis et al., in a lightning talk at Evolution, take advantage of these new genome-sequencing techniques to approach the genetics of adaptive radiation in Anolis. To understand the genetic mechanisms underlying the adaptive radiation of anoles, they preformed de novo genome sequencing on three Anolis species (Anolis frenatus, Anolis apletophallus, and Anolis auratus), chosen to capture different sub-groups of the Anolis phylogeny. With these data, and the published genome sequence of Anolis carolinensis, they looked for patterns in the rate of evolution compared to other vertebrate groups. They also looked within the Anolis genome to detect specific genetic regions associated with selection across the anole radiation.

Tollis et al. found that, in general, anoles appear to have a high rate of molecular evolution for a vertebrate species, which may parallel the high rate of phenotypic evolution seen in this clade. In addition, Tollis et al. looked for signatures of selection across the four Anolis genomes and identified regions associated with reproduction, olfactory reception, and limb development. This last category is of special interest, given that anoles are notorious for changes in limb morphology between species and that limb morphology is one of the key components of ectomorphs in the Greater Antilles. Tollis et al. have provided a great example of using new genetic tools to approach fundamental questions about the mechanisms underlying adaptive radiation.

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