Anole Research In Colombia And Venezuela: Scientist At Work Blog Posts

Earlier this spring, I reported on a trip to Colombia and Venezuela to collect natural history data on several unknown anoles: A. heterodermus, A. onca, A. annectens and A. tigrinus. As part of the Scientist at Work: Notes from the Field blog on the Science webpage of the New York Times, I wrote a series of posts, which have now all been published. If you’re interested, links to them are provided below.

Losos, J.B. March 12, 2013. Quest for a little-known suburban lizard. link to article

Losos, J.B. March 14, 2013. Life of a Lizard Wrangler. link to article

Losos, J.B. March 19, 2013. Lizard Olympics. link to article

Losos, J.B. April 2, 2013. A Beach Vacation, Lizard Style. link to article

Losos, J.B. April 11, 2013. An Embarrassment of Anoles. link to article

Losos, J.B. April 25, 2013. A Night of Saurian Spotlighting. link to article

There Is Always Something New To Learn: Lizard Eggs Can Survive Flooding

A close-up of the hatchling, un-hatched egg, and the remains of the hatched egg, we recently reported on.

A close-up of the hatchling, un-hatched egg, and the remains of the hatched egg, we recently reported on.

It is interesting that even in fairly well-studied species, we can still find new information from chance observations and carefully planned empirical studies. Recent studies have found that brown anole (Anolis sagrei) females favor nest sites that are quite moist. Losos et al. (2003) also found that the eggs of this species can survive being inundated for up to six hours by seawater. Recently we reported on an observation of brown anole eggs that remained viable after being inundated by fresh water due to heavy rains. The difference between the two conditions is, since seawater is more saline than the content of the eggs, it can be expected that the eggs would lose water, while when exposed to excessive fresh water the water gains will exceed losses, and the eggs will swell.

The observations of other researchers and ours suggest that the eggs of A. sagrei are fairly tolerant to a wide variety of environmental conditions and can remain viable when exposed (for a few hours) to excessively wet conditions. Our observations in Taiwan also reveal that unless washed away or damaged, the eggs of A. sagrei can remain viable during the high rainfall (often as a result of typhoons) of this region.

As Aristotle (384 BC – 322 BC) said, “In all things of nature there is something of the marvelous.” And I would like to encourage everyone to look into their chance observations and to describe them, not only in chat forums, but also in journals as natural history notes. That way the information is not only disseminated, but may also be accessible to a greater audience and for generations to come.

Brazilian Samba Giant And Anole Biologist Paulo Vanzolini Passes Away At 89

The doyenne of Brazilian herpetology, Paulo Vanzolini died two days ago at an age of 89. Renowned for his herpetological expertise, Vanzolini was even more famous for his samba compositions. Indeed, his Wikipedia page focuses more on his music career and discography than his zoological contributions, an oversight that perhaps Wikipedia-savvy AA readers can rectify.

 

With regard to anoles, Vanzolini made two important contributions in collaboration with Ernest Williams. The first was their monographic treatment of variation in the mostly-Amazonian Anolis chrysolepis species complex, a group that was recently revised by D’Angiolella et al. This monograph was notable not only for its detailed study of geographic variation, but also for its suggestion that speciation may be prompted by climatic cycles that lead to contraction of populations into isolated, allopatric pockets of suitable habitat. This hypothesis was suggested simultaneously and independently of the same suggestion for Amazonian birds by Haffer and led to the influential and much-debated Pleistocene Refugia hypothesis, discussed previously in AA‘s pages.

Vanzolini and Williams had a follow-up paper that is less well-known, but equally insightful and ahead of its time. In this paper, the authors argued that such refuges might be particularly important foci of adaptive evolution and speciation when they disappear entirely. The argument is that populations might be trapped in refuges and as the habitat continues to deteriorate over time, the populations might have no choice but to adapt or perish. In this way, arguing from the chrysolepis complex as well as from other lizards, dry forest or even grassland species might evolve from wet forest ancestors. This hypothesis could explain the existence of closely-related species occupying very different habitats–the antithesis of what is now known as phylogenetic niche conservatism–and even might explain the parapatric distribution of close relatives if the new species expanded its range (see previous post for more discussion). This idea was published in Papéis Avulsos de Zoologia in 1981 .(download it here) and deserves more attention than it has received.

Evolutionary Diversification Of Caribbean Snakes

The Puerto Rican Racer, Alsophis portoricensis. Photo by Donald Gudehus

Sometimes it’s easy to forget that anoles aren’t the only animals in the Caribbean. But, in fact, there are other types, even of reptiles, and some of them have diversified a fair bit (though none, of course, to the extent of anoles). One such group are the alsophiine snakes, formerly all in the genus Alsophis. This Caribbean radiation of racer-like snakes includes at least 43 species ranging in size from 200-2000 mm in length and occupying a variety of habitats.

burbrink et alRecently, Frank Burbrink and colleagues, in a paper in  the Journal of Biogeography, have re-analyzed DNA data originally presented by Hedges et al. and have investigated rates of species, morphological and ecological diversification. The phylogenetic tree they recover is very similar to the Hedges et al. phylogeny and indicates fairly extensive within-island diversification. Sounds very anole-like, but it turns out that rate of diversification is quite different. Unlike anoles, species diversification and the evolution of morphological variety putter along a fairly constant rate (with a few statistical twists and turns).

Why the difference? Burbrink et al. postulate that the opportunity for diversification has been just as great for alsophiines as for anoles, so why are the evolutionary patterns different? The authors put forward a number of possible explanations, but none is compelling. Of course, although adaptive radiations often exhibit explosive bursts of diversification, there is no necessity for this to occur, and some very diverse groups have radiated at a more sedate pace. Moreover, one might question why alsophiines haven’t diversified even more–sure, they differ in body size and climatic niche, but how different are they otherwise? And how many species can co-occur at a given locality? Is it just lack of time–one of Burbrink et al.’s hypotheses–or is something constraining alsophiine diversification?

More generally, it would be interesting to conduct similar analyses on other Caribbean taxa–not just reptiles, but also amphibians, birds, even insects and plants–to see what generalities, if any, characterize Caribbean evolutionary diversification.

Weekend Brown Anoles From Daffodil

Photojournalist, anole aficionado and AA has devoted yesterday’s post to the goings-on of her local brown anoles. Check it out Daffodil’s Photo Blog.

True Facts About Chameleons

httpv://www.youtube.com/watch?v=UR_byRbXxvs

Anole researcher Anthony Herrel showed his true colors recently in a seminar when he said that chameleons are cooler than anoles. Be that as it may, this video is a fun demonstration of the amazing traits that make chameleons the second coolest lizards. It’s part of the “True Facts” series of videos, which are quite entertaining–one episode on tarsiers was featured in a recent AA post.

Katydid Eats Anole

princeps eating by conehead katydidHere’s a disturbing photo that came across Facebook the other day. And I always thought these katydids seemed like delightful, gentle forest nymphs. Who knew they could be vicious killers. This comes from The Biodiversity Group’s Facebook page, but I couldn’t locate the image on their website, though I didn’t look that hard. They identify the anole as A. princeps but don’t provide any further information.

More Surprises From Anolis Tigrinus

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Anolis tigrinus. Photo by J. Losos.

I’ve previously posted on the surprising behavior of the Venezuelan A. tigrinus. We found them in astonishing abundance in the town of Colonia Tovar, in the mountains above Caracas. Despite their twig anole appearance and their twig anole habitat use–on twigs and other narrow surfaces–they don’t behave like twig anoles. Rather, they move rapidly and often, and seem to display a lot as well. Some Caribbean twig anoles move frequently, albeit slowly, and some display a lot, but none zip around like these guys. Moreover, we’ve seen a number of other mainland twig anoles, and they live life in the slow, slow, slow lane. So, our first day of tigrinus-watching was full of surprises. But that night, things got even more surprising, as I recently recounted in my most recent post in the Scientist at Work blog of the New York Times..

The tell-tale banded tail.

The tell-tale banded tail.

Anolis tigrinus sleeping on a stem. Photo by J. Losos.

Anolis tigrinus sleeping on a stem. Photo by J. Losos.

As we walked down to the local brewhaus to grab a pizza, we casually scanned our surroundings. And as we walked by a weed patch, there was the tell-tale white blob hanging on to the end of a grass blade. It could be only one thing–an anole. In the Caribbean, this would be primo grass-bush anole habitat, but there were no grass-bush anoles here. Quick examination confirmed that it was an A. tigrinus. That’s right, a twig anole in the grass. And then as we looked around, there were more and more of them–the lot was silly with twig anoles.

The next two days, I came back during the day to look for twig anoles. Unlike in the woods where we had found them initially, the twig anoles here were hard to find during the day, even though the place was full of them at night. The few I saw were deep in the vegetation–my conclusion is that this guys are moving through the vegetation, navigating along narrow branches, stems, and grass-blades.

Which leads, of course, to the question of why in the Caribbean, grass-bush anoles use low-lying narrow vegetation in grassy, bushy  areas, while twig anoles use narrow twigs and branches up in the trees. Perhaps A. tigrinus  is just being opportunistic, taking advantage of an unoccupied habitat and moving in? Certainly a plausible explanation, but I’ve never, ever heard of a twig anole in the grass in the Caribbean. Has anyone? Not even in Jamaica, where there are no grass-bush anoles.

This in turn reminds me of the trunk-crown anoles, A. allisoni, I observed in the grass in Roatan, Honduras. If nothing else, it’s important to remember that anoles are very behaviorally flexible and adaptable. Even though they’ve specialized to different microhabitats, they aren’t so specialized that they can’t use other microhabitats when they get the chance. Crown-giant anoles on the ground are one example, but that’s another story.

In any case, A. tigrinus is a very interesting anole, and it’s great abundance would make it an excellent choice for behavioral and ecological studies.

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Sweet dreams, little tiger anole. Photo by J. Losos

Nothing Worse Than Being Stuck In The Rain When A Predator’s Around

Anole perch height depending on whether it was raining and whether curly-tailed lizards (Leiocephalus carinatus) were observed on the plot.

Do you like standing out in the rain, especially when it’s cold? Me, neither. But that’s what the dastardly curly-tailed lizard forces brown anoles to do. Any sensible, semi-arboreal lizard would come down from the heights and seek shelter when it starts to rain, and that’s exactly what brown anoles do. Except when they’re in areas of high curly-tailed lizard activity, in which case they suck up and stay up high, shivering and being pelted by rain drops. That’s what research by Marta Lopez-Darias and colleagues (among which, yours truly) reported in a recent paper in Ecology. As the figure below illustrates, pretty much the only time the brown anoles drop down is when the weather goes to pot and curlies aren’t around: cool, windy, and very humid–in other words, when it’s raining. But if big boys have been cruising around on the ground, the anoles maintain their high perches.

Brown anole perch height as a function of a variety of weather variables and of curly-tailed lizard activity (in this figure, instead of presence/absence as in the figure above, predator activity was measured as the time-standardized number of active curly-tailed lizards observed on the plot).

All kidding aside, it’s not clear why they come down when it’s raining, but presumably there’s a benefit to it. One can only speculate what that is; my first guess: when it’s wet and cold, anoles are less able to notice approaching predators and less able to get away quickly because of their lower body temperature, hence they seek safer environs. Or perhaps there’s simply no potential prey afoot, and thus no reason to hang out in a high vantage point looking for them. Whatever the reason for doing so, it appears to be overruled by the threat of marauding curly tails.

As for details of the study: ten study plots were set up in various parts of Great Abaco. Plots were regularly censused, tabulating the number of curly-tailed lizards observed, the perch position of each anole observed, and a battery of meteorological variables.

There Is More To That Beach Anole

Norops-onca-male

Adult male Anolis onca from Isla de Margarita basking.

Continuing the recent interesting post on the Beach Anole Anolis onca, I decided to write something about my personal experiences with this amazing species and attempt to summarize some of what already exists in the literature. Famous by its lack of expanded, smooth, infradigital lamellae, there is a lot more to these beautiful lizard.

Coastal-thorn-scrubland

Typical habitat of Anolis onca on Isla de Margarita

This “beach” anole is basically endemic to Venezuela (it is also found on a narrow portion of adjacent Colombia). Within Venezuela, Anolis onca has a disjunct distribution (more on that below). It ranges continuously along the coast of the states of Zulia and Falcon in the West. It is also found on the islands of La Tortuga and Margarita, as well as along the coast of the state of Sucre in the East. Another, possibly disjunct population has surprisingly been recorded from dry savannas well inland in the state of Monagas! I have observed A. onca in western Venezuela, but I am most familiar with populations from Isla de Margarita. I have traveled countless times to the island and since I was a kid I always remember being fascinated by these fairly large, active anoles. On Isla de Margarita, A. onca is definitely ubiquitous. It is easily found on thorn scrubland, coastal sand-dune environments, and beaches. It is also common around human habitation.  I have always observed this anole in sites with constant and strong wind currents. Several authors have suggested that the windy conditions present in the habitats preferred by this lizard may have prevented it from being strictly arboreal like other anoles (Williams, 1974; Miyata, 1975; Kiester, 1977).

onca-dewlap

These guys have a really large, beautiful dewlap

Whatever the case, it is definitely more terrestrial than other anoles (even species that are commonly found on the ground, e.g. Anolis planiceps) and I have observed it numerous occasions on open ground. However, it often climbs up to about 1.5 m on shrubs, cacti, vines or rocks. Around human habitation it also perches on unfinished walls of buildings, cobble and fence posts (same places frequented by the larger Tropidurus hispidus on the island). I also have observed A. onca on open sand banks of beaches in close proximity to salt water (Ugueto and Rivas, 2010). Williams (1974) also mentioned finding this species near the seashore. Interestingly, light colored specimens are very well camouflaged amidst the sandy soil. I have noticed that when A. onca perches on low shrubbery it often just exposes its head above the leaves. I do not recall seeing such behavior in other anole species.

Most individuals remain motionless when first spotted. If you get too close for comfort, lizards invariably run towards and hide within nearby bushes, clumps of herbaceous vegetation or thorny shrubs. Occasionally they may hide beneath rocks or boulders. Collins (1971) reported that some specimens escape into Ocypode crab holes after lizards were pursued for long time, but I have never observed this behavior. Various types of small arthropods like grasshoppers, robber flies, beetles and spiders have been reported as prey (Roze, 1964; Kiester, 1977; Ugueto and Rivas, 2010). Kiester (1977) reported that the analysis of 38 stomachs revealed that a particular species of chrysomelid beetle constituted a large portion of the diet in western Venezuelan lizards. Saurophagy has also been reported in this species; Miyata (1975) and Kiester (1977) recorded a large individual preying on a female Cnemidophorus lizard in northwestern Venezuela. I observed this species preying on small flies that passed near the lizard on a beach on Isla de Margarita. Kiester (1977), however described a very interesting mode of predation in which the lizard stalked fast moving robber flies in a cat-like fashion using the irregularities of the terrain as cover and dashing towards prey when at close range. What is known about the reproduction of this species is fragmentary at best, but it appears it could be seasonal on northwestern Venezuela. Kiester (1977) reported females laid eggs after the rainy season so that hatchlings come out during January, February and probably March.

Anolis-onca-map

Distribution of Anolis onca and the closely allied Anolis annectens in Venezuela. Notice the disjunct distribution of onca.

As I mentioned earlier, the distribution of A. onca in Venezuela is disjunct and the western and eastern population are separated by a long stretch of apparently suitable habitat.

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