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

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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.

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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).

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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.

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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.

World Turned Upside Down

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Knight anoleYour intrepid correspond is currently in West Palm Beach, Florida, about to embark on a meandering trip north in quest of yellow-dewlapped brown anoles and other anole curiosities (speaking of which, anyone know a good site to find A. sagrei with a lot of yellow in their dewlaps? Like half or more?). In any case, today’s curiosity occurred as I was walking through a large condo development. Out of the corner of my eye, I saw something double surprising. The first surprise was that it was a knight anole–I wasn’t aware that they had spread so far from Miami, although a subsequent google search revealed that, in fact, they are not only known from West Palm, but also from considerably further to the north (see map above).

The second surprise was where the lizard was. It was on the ground. Well, from the corner of my eye, it seemed like it had hopped from the ground, but when I actually turned to look, it was on a tree trunk, about 10 cm off the ground. Still, quite low, not where you expect to find a crown-giant. Sean Giery coincidentally wrote a post about female knight anoles laying their eggs in holes on the ground, so that’s a possible explanation, though this was a little knight, more of a knightlet, at ca. 130 mm svl.

IMG_2120xIMG_2119xExcited by the spotting of such a noble beast, I forsook the brown anoles and began looking for the big greens. And they were remarkably abundant! In about an hour of meandering, I found 13! Some were high up in the tree, right where they should be (you can see the silhouette of one in the photo to the left), but others were much lower (photo on right), and one was on a shuffleboard court (below)! In addition, as I posted recently, evidence of terrestrial habitat use was also found post-mortem by the discovery of a two-dimensional equestris on the one-lane road running through the complex.

Site of the terrestrial knight anole spotting. A careful look will reveal that Anolis equestris isn't the only terrestrial lizard in this habitat.

Site of the terrestrial knight anole spotting. A careful look will reveal that Anolis equestris isn’t the only terrestrial lizard in this habitat.

IMG_2101Why is it that these lizards are on the ground? Well, for one thing, this is not a forest, but a condo subdivsion. There are plenty of trees, but most have been planted and they are spaced out. Many of the knight anoles I found were in trees there were completely isolated, such as the one in the photo on the left.  In other words, to get from one tree to another, or to colonize a tree in the first place, the big lizards have to move on the ground. There’s been very little work on knight anole ecology and habitat use–it would be really interesting to see how often they move from one isolated tree to another.

IMG_2117But these low-riding knights (in the Jansonian sense) are only half of the reason that I’ve titled this post “World Turned Upside Down.”

Anoles Feeding On Liquids – Please Help

Anole feeding on nectar. What about sap? Photo by Sparky Leigh.

Good day everyone. I am currently working on a short manuscript about a brown anole that I observed feeding on sap. I am aware that some anoles will feed on nectar (see list below), but I would like to know if anyone has ever observed anoles feeding on sap? If you have any references pertaining to anoles feeding on nectar or sap, and it is not listed below, would you please e-mail it to me at a_sagrei@hotmail.com, or at least provide me with the reference so that I can try to obtain it myself. Thank you very much.

Gerrut

P.S. Lizards rule!

References I am aware of:

Campbell, T. and C. Bleazy. 2000. Natural history notes:  Anolis carolinensis (green anole). Nectivory and flower pollination. Herpetological Review 31: 239.

Colón Archilla A.D. 2010. Nectivory in Puerto Rican emerald anoles (Anolis evermanni). IRCF Reptiles and Amphibians 17: 144–145

Echternacht, A.C. and G.P. Gerber. 2000. Anolis conspersus (Grand Cayman Blue throated Anole). Nectivory. Herpetological Review 31:173.

Liner, E.A. 1996. Natural history notes: Anolis carolinensis carolinensis (green anole). Nectar feeding. Herpetological Review 27: 78.

Okochi, I., M. Yoshimura, T. Abe, and H. Suzuki. 2006. High population densities of an exotic lizard, Anolis carolinensis and its possible role as a pollinator in the Ogasawara Islands. Bulletin of the Forestry and Forest Products Research Institute 5: 265–269.

Perry, G. and J. Lazell. 1997. Anolis stratulus (Saddled Anole). Nectivory. Herpetological Review 28:150–151.

Perry, G. and J. Lazell. 2006. Anolis pulchellus (Grass Anole). Nectivory. Herpetological Review 37:218–219.

Rios-Lopez, N. 2004. Anolis stratulus (Saddled Anole). Extrafloral herbivory. Herpetological Review 35:386.

Timmermann, A., B. Dalsgaard, J.M. Olesen, L.H. Andersen, and A.M. Martin Gonzalez. 2008. Anolis aeneus (Grenadian Bush Anole). Anolis richardii (Grenadian Tree Anole). Nectivory/pollination. Herpetological Review 39:84–85.

Valido, A.M. 2006. Anolis allisoni (Allison’s Anole/Cameleon Azul). Nectar feeding. Herpetological Review 37:461.

Evo-Devo Meets Sexual Dimorphism: The Face Does Not Tell The Whole Story

Males and females of many species vary in their morphology, behavior, and physiology. Whether exaggerated weapons, elaborate coloration patterns, or dramatic differences in size, these sexual dimorphisms form some of the most eye-catching elements of biological diversity. These striking differences are often considered as a product of sexual selection, whether due to direct female choice for an elaborate structure or traits used by combative males to assert dominance. But additional patterns of dimorphism become visible with quantitative comparisons of male and female body proportions, which may yield additional clues to ecological differences between the sexes. Considering that patterns of sexual dimorphism can diverge rapidly among species it is no surprise that they have intrigued biologists since before Darwin.

Anolis brunneus from Crooked Island, Bahamas. A member of the carolinensis clade of anoles exhibiting extreme levels of facial length dimorphism.The male is the large lizard to the bottom left, the female to the upper right.

Nesting Knight Anoles

Female Knight anole digging nest at the base of an oak.  April 2013

Female Knight anole digging nest at the base of an oak. April 2013

On my way to teach biology lab the other day, I ran into this female knight anole (actually, my buddy Zack is the one who spotted it).  She was obviously spooked by us and after snapping a few photos I backed off to see if she would continue nesting.  After a few seconds she returned to her task.  I don’t know why I imagined anoles would dig with their hind legs, but for some reason I did. However, she continued to excavate with alternating strokes of her front feet.  Unfortunately I had to run to a meeting, but when I returned a few hours later her hole was still there.  Zack had stayed behind to watch her progress and reported that she had aborted the endeavor when a bicyclist whizzed past a bit too close. The hole was about 4 centimeters deep and 5 wide into the mineral soil when she left.

Female Knight anole observed digging nest at the base of a large fig tree.  Note the soil under her front claws and on her snout.  Observed July 2012.

Female Knight anole observed digging nest at the base of a large fig tree. Note the soil under her front claws and on her snout. Observed July 2012.

This is the second time I’ve seen nesting knight anoles in Miami.  The first was last summer when I found a very healthy looking female at the base of a fig tree.  There was dirt on her snout and a small hole where she had been.  In both cases, the nesting females were within 1 meter of a tree, and both were excavating with their front feet and possibly their snout.  Also, the dates of each observation show that there is a protracted nesting season in South Florida that includes April 10th to July 20th.  Such a long season explains the variable size of first year knight anoles that I find during the spring.

Anolis Equestris Dead On Road

equestris dor2xHere’s something you don’t see every day. More on these guys soon.

Anolis Annectens, The Retrograde Anole

Anolis annectens. Photo by J. Losos.
Anolis annectens. Photo by J. Losos.

No flies on you guys–this remarkably obscure anole was quickly identified. So, what’s its claim to fame?

Well,  we have to backtrack to Anolis onca for a minute. Faithful AA readers will recall that A. onca is the only anole lacking a subdigital toepad. For this reason, at one point it was placed in its own genus, Tropidodactylus. However, in 1974 Ernest Williams described a new species in Breviora based on a single specimen found in a jar of A. onca in the Field Museum in Chicago. At that time, Anolis was characterized by having expanded scales under phalanxes ii and iii of the toe, whereas Tropidodactylus, of course, had none (phalanxes are the bones in a toe–humans have three per finger, for example). What was remarkable was that the new specimen had expanded scales under phalanx ii, the Anolis condition, but only keeled scales under phalanx iii–it was intermediate between the two, hence the specific name annectens. Here’s an image from Williams’ paper and a photograph of an annectens toe.

From Williams (1974).

From Williams (1974).

Anolis annectens foot. Photo by J. Losos.

Anolis annectens foot. Note the lamellae under only one toe bone. Photo by J. Losos.

 

 

 

 

 

 

 

 

The term “retrograde” comes from Williams’ paper and refers to the idea that A. annectens and A. onca illustrate a morphocline in toepad reduction–halfway gone, then all the way gone. Incidentally, recent molecular studies confirm that the two species are sister taxa.

Amazingly enough, A. annectens occurs in the same general region as A. onca, near Lake Maracaibo in Venezuela. In 2007, Tito Barros and colleagues reported in Tropical Zoology on the collection of an additional 22 specimens of A. annectens, confirming that the one individual wasn’t a freak. They also provided information on coloration, geographic distribution (it still hasn’t been found in sympatry with A. onca) and some data on ecology.

Since we were in the area any way, we decided to go and look for them after collecting data on A. onca. By “we,” I mean Tito Barros, Gilson Rivas, several students of theirs, and Rosario Castañeda. Anthony Herrel was busy back at the field lab conducting performance trials (sprint speed, bite force), and I was on assignment finding discarded cardboard boxes to make a proper racetrack.

The weather was beastly hot, about 110 degrees Fahrenheit, with no wind. Here’s what Rosario had to say:

Anolis conspersus, UV Dewlap Photos And Anoles As House Geckos

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On a recent trip to Grand Cayman I was interested in the UV reflecting dewlap of Anolis conspersus. The dewlaps of these lizards appear blue to our visual system but are maximally reflective in the ultraviolet. While anoles have 4 cone types (ultraviolet, blue, green and red sensitive), humans have only 3 and cannot see UV light so to understand what these lizards look like in the UV, we have to use specialized camera equipment.  The photo to the right shows what a displaying A. conspersus looks like to our camera system when imaged in the human visual spectrum as commercially available digital cameras also have only three channels corresponding to the three human cone types.  Presumably if we were also able to see in the ultraviolet as many other animals can, our cameras would be designed with a separate channel for ultraviolet.

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These images of the lizard in the UV show clearly the regions of the dewlap and that are highly UV reflective and the pattern of UV reflectance in other areas.  One somewhat interesting finding is that while the dewlap scales are highly reflective across the human visual spectrum (which is why they appear white to our eyes) they reflect very little UV light.  The lower photo is a monochromatic image (both the red and blue channels in this camera are sensitive to UV so the raw image appears purple) that makes it a bit easier to see brighter areas as white.  Note how bright the dewlap appears relative to the reflectance standard, when imaged in the human visual spectrum a similar monochromatic image of the dewlap would appear very dark.  I believe this shows the potential value of UV photography when studying Anolis dewlap patterns.  While the UV nature of the A. conspersus dewlap is uniform, it’s likely that other species have patterns visible in the UV we’ve previously missed.  We have also used this UV photography setup in SE Asia to image Draco flying lizards and other species, some of which have patterns that are visible only in the UV band.  The goal of this project is to make a camera system with pixel channels similar to the four cone types found in Anolis lizards and birds to image whole organisms and really “see” the patterns organisms experience with their visual system as they would see them.  As Anolis visual pigments and their associated oil droplets appear to be fairly conserved, this seems to be achievable.

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Another surprise (to me) was the large number of A. conspersus on Grand Cayman using lights at night to feed.  I’ve spent many months doing fieldwork in SE Asia and Central America and can’t recall seeing this sort of thing with other diurnal lizard species, but on Grand Cayman it was quite common in A. conspersus.  I observed one A. conspersus male chase away a Hemidactylus that got too close to the light, showing that the anoles at least occasionally displaced the group I typically associate with feeding around lights.  A check of the literature shows this has occasionally been documented on other Caribbean islands, but as far as I can tell no one has published on this in mainland species.  What diurnal lizard species have others observed using lights to feed at night?

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