Photo-chronicler of Floridian natural history Karen Cusick has done it again. We’ve been captivated by her backyard photos before, but here’s photo of a female green anole with sand on its snout. Been digging holes to bury her eggs, maybe? And while Karen observed the little lady lizard, it suddenly darted into the bushed and emerged with a meal!
Lizards are active creatures, often running around in new territories to explore and find food. Sometimes they encounter challenges that keep them from running too far. When they wander away from home, how do they get back? It’s a question that’s led researchers to study this topic.
After watching the daily routines of Anolis lizards by using tracking devices placed on their backs, researcher Manuel Leal learned that they return to the same home again and again. This established the next question which was to find out how the lizards knew how to get back. Birds have a similar ability to find their way home. Although the exact method has not been discovered, it’s possible that lizards have similar abilities and functioning as birds in finding their own again.
They Claim Their Homes
Anolis lizards, especially males, claim trees as home territories, fighting to keep any newcomers off their bit of land. They’ve proven that they remember exactly where they stake out their claim, and like all animals, they like structure in their environment, including the location of where they spend their days and nights. Some studies prove that after disorienting the lizards and placing them a far distance from their home, they can still find their way back within 24 hours.
Then They Listen
U.S. Geological Society geologist John Hagstrum proposed that in order to get back home, pigeons use sounds wave; extensive studies on pigeons show that they use low-frequency sound waves to create an acoustic map of where they are. This way, they can identify predators and safe spaces to land. Some have wondered whether Anolis lizards might have similar capabilities that are advantageous for homing.
I’ve taken more than four hundred toepad pictures using the new macro photography technique I introduced in an earlier post and I’ve learned a few tricks that I want to share in this update.
First and foremost, I highly recommend this approach. For those of you looking to capture a lot of toepad data, particularly in the field, this kit is way faster and more portable than using a flatbed scanner and the images I’m getting are at least as sharp.
A few tips:
Petri dishes work great as a clear platform to place the lizard feet on. I found that the 60 mm diameter dishes were much easier to balance atop the lens (~40 mm in diameter) than the larger dishes I’d originally shown.
I cut and taped a scale bar to one edge of the petri dish so I wouldn’t have to worry about juggling a lizard and a tape measure.
Make sure you have several petri dishes – they scratch fast – and keep some ethanol and a kimwipe close at hand.
The app that lets you remotely trigger your iPhone is absolutely maddening. Do not download it. I’m not even going to relink the name. Instead, I suggest a much more stable alternative: connect your phone to your computer with the USB cable, open QuickTime Player, select File > New Movie Recording and click the down arrow next to the record button. This will give you the option to select your attached iPhone as a recording device. This live-view is far more stable and less frustrating. *Windows and android users I’m afraid I haven’t had an opportunity to sort out a solution for those platforms. If you know of something that works, please include in the comments!
Unfortunately, through the live view all you can see is whether the lizard is in position. You cannot remotely trigger the shutter this way. That means you’ll need a second pair of hands to help. I found it worked best when my partner was in charge of putting the ID tag in the frame after I’d placed the lizard foot and then pushing the volume button on the side of the phone to trigger the camera shutter.
Lighting is really important. I suggested a headlamp in the previous post providing an oblique light source through the diffuser around the lens. I tried using a microscope fiber optic light source but I was really unhappy with the “warmth” of the light. I found that the white-LEDs in my headlamp produced a much more realistic looking image (see above). Also, make sure you don’t have any light sources above/behind the subject. Backlighting confuses the camera’s auto-contrasting and results in dark and sometimes unfocused images.
After scampering about much of North America the past few decades, I once again live in my hometown of Ormond Beach, Florida — on the northern edge of Volusia county. When I was a kid, back in the late 70s and early 80s, I spent much of my time tangling with and studying our local anoles. The Carolina greens (A. carolinensis) were dominant back then, covering our walls, windows, trees, and (sometimes by forced measure) our ear lobes. Every now and then I’d find a Cuban brown (A. sagrei) — usually around the shopping centers and strip malls. Nowadays, of course, that coin has flipped. The Carolina greens have moved back up into the higher foliage and the Cuban browns dominate our shrubs, walls, and windows.
I remember actually finding a Cuban brown anole on our property in 1984 or so. I was in 4th grade, drunk on Star Wars and lizards. I managed to catch the little non-native lizard and put it in my anole terrarium (a homemade wood-and-open-screen enclosure my dad and I built). I was in the habit of catching anoles (and the occasional snake), keeping and watching them for a day or two, and then releasing them back into the yard. Needless to say, the Carolina green already in the enclosure wasn’t too thrilled with his new roommate. Though guilt eventually kicked in the following day, I admit I was somewhat delighted by the defensive/discomfort color play of that poor Carolina green. Usually, they’d be cool, smooth emerald green with very little patterning… but distressed or riled up Carolina greens certainly know how to put on a good color and pattern show.
Soon enough, I released the Carolina green back into the yard and kept the Cuban brown for another day or two. This little moment of tension, however, leads me to the point of this post: the distress patterns of our local Carolina green anoles. More specifically, I’m interested in the presence of a supraspacular dark spot that shows up with some individuals. It’s a dark spot with light trim that sometimes appears just above and behind the front shoulder line — as seen in this particularly ornate individual photographed in Miami-Dade county on 18 March 2017:
This Miami-Dade individual really stuck out to me. It’s patterning was distinct. It was quite large. It had that supraspacular spot. Most notably, it was still wielding quite a bit of green. Could this be A. porcatus? Like many naturalist-lizard enthusiasts, I tend to catch myself up in the eternal cycle of porcatus-or-not? when I’m in south Florida. Heh. Nowadays, my assumptions generally fall on the side of A. carolinensis unless I’m with somebody more in-the-know who can tell me differently with confidence; this hasn’t happened yet. Honestly, I have a hard time seeing a clear difference between the two. I’m glad I’m not alone.
Though distinct, this fabulously mottled Green wasn’t the only Green I’ve photographed with that supraspacular spot. Here’s an impressive male tangling with a Cuban brown anole in the Lower Keys of Monroe county, Florida, on 08 June 2007:
Further north, in my home territory, I’ve only noticed and photographed two individuals with that spot, albeit with less figure-ground contrast between the spot and the trim.
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 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.
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, 2, 3), 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.
Red-legged wandering spider (Cupiennius coccineus) consuming a house gecko (Hemidactylus frenatus) at Sirena Biological Station, Corcovado, Costa Rica
When someone first asked me about the major predators of anoles, my first thought was to talk about curly-tailed lizards (Leiocephalus carinatus) in the Caribbean, vine snakes (Oxybelis spp.) in the neotropics [see my previous post on anole predation by O. aeneus at La Selva], and birds. I think that as herpetologists, we tend to fall into the trap of thinking of invertebrates as “lesser” taxa to be preyed upon by small vertebrates like lizards, and in turn for small vertebrates to be eaten by larger vertebrates.
I, too, when thinking about how selective pressures shape morphological variation in mainland and island habitats turned to fellow herps and birds as the primary predation pressure for mainland anoles. However, it wasn’t until I arrived in Costa Rica that I discovered the high prevalence of voracious arthropods, and I realized that our beloved lizards had much more to fear!
Orange wandering spider (Cupiennius getazi) with egg sac at La Selva Biological Station, Costa Rica
Red-legged wandering spider (Cupiennius coccineus) eating a pink katydid (Tettigoniidae: Phaneropterinae) at La Selva Biological Station, Costa Rica
A large adult female mantis (Phasmomantis championi) at La Selva Biological Station, Costa Rica
Conehead katydid (Tettigoniidae: Conocephalinae: Vestria sp.) at La Tarde, Osa Peninsula, Costa Rica
In a single night at La Selva, I could easily find dozens of large wandering spiders (Ctenidae), and if I pointed my headlamp higher in the trees I could see eyeshine from hundreds of spiders. Given the high density of large ctenids at La Selva, it is not unlikely that anoles and small tree frogs constitute a major portion of their diet. In fact, I wouldn’t be surprised if large arthropods are one of the most common predators of mainland anoles in some regions.
The same might be the case for giant mantids of the genera Macromantis and Phasmomantis, and conocephaline katydids sporting fearsome mandibles (e.g. Copiphora spp.). Since the invasive Chinese mantids (Tenodera sinensis) in North America are well documented to prey on hummingbirds almost equal in size to the mantids [see Nyffeler et al. 2017], surely larger and bulkier species in the neotropics can take lizards much smaller than themselves. Even though wandering spiders and conehead katydids are primarily nocturnal hunters, I have heard many stories of these arthropods being implicated in anole and tree frog predation. Research looking into how ctenids and nocturnal katydids forage would help determine if they can actually detect sleeping anoles or if predation events occur from the arthropods simply running in to the anoles while on the move.
If anyone here on Anole Annals has any anecdotal or photographic records, please comment below.
To throw a twist on this discussion, is it possible for a spider to prey on a lizard two and a half times its size? A new paper about a vertebrate-eating jumping spider (Salticidae) describes just that! Considering arthropods as possible major players in anole predation could shed light on behavioral and ecological studies of mainland anoles.
Figure 1 from Nyfeller et al. 2017, showing female jumping spiders (Phidippus regius) consuming Carolina anoles (Anolis carolinensis) and Cuban tree frogs (Osteopilus septentrionalis)
Here are a few more spider photos to wrap up this blog post.
Getting good pictures of lizard toepads in the field can be tricky. Flatbed scanners are heavy and don’t take well to transit bumps and bruises, and getting a digital camera to focus on the toe, not the glass, requires surgical precision on the manual focus ring. I’ve just found a new solution for an iPhone (or GooglePixel, if that’s how you roll), and I’m eager to share.
Here’s the setup in action (and, by the way, this particular lizard’s bite force was classified as medium-ouch):
You’ll notice that when the camera is facing up the iPhone screen is facing down. Obviously this makes it difficult to snap the photo—enter the app WiFiCam. This app enables you to type the phone’s IP address into your web browser and remotely trigger the camera, as long as both devices are on the same wifi. It’s very simple, and the price was right (free!).
And so here’s the whole shebang:
(Don’t forget to keep a tissue handy for wiping up lizard poop!)
And not to bury the lede, but the results are fantastic (see above).
A few things to note:
The white plastic platform around the lens ensures perfect focal distance so getting your lizard as close to that plane as possible is ideal. I tried a square of single pane glass but wasn’t tremendously pleased with the results. The above is taken with a cheap plastic petri dish, which works great but scratches quickly. Another option I’m going to look into is a glass microscope slide. (The biggest drawback to the slide is that it’s smaller than the camera lens platform… meaning that the lizard can actually poop ON YOUR PHONE. And believe me, they will.)
The app works fine for controlling the shutter, but it’d be nice to be able to also control other camera settings like focus point and brightness or contrast. There might be other apps out there that do all of that; I just haven’t tried to find them yet. If you’re taking photos of lizard toepads in a place without wifi (as you most likely are), you can use your computer to create a local network and pair the camera to the computer that way.
I found that the sidelight was really helpful to get good illumination on the toes. Without the sidelight the camera sometimes adjusts for ambient light behind the foot, making the lamellae hard to see. My headlamp was the perfect size and brightness and worked great.
One last thought: Moment also has a fisheye lens that might do a really nice job of canopy cover photos in the field. That’s on my short list of things to experiment with in the near future!
I’d love to hear your thoughts on how to improve the system in the comments.
I recently saw these four anoles on a trip to Costa Rica. All four were sighted in La Fortuna in the province of Alajuela. They were sighted on July 18th and 19th, all within a few meters of a forested stream. I have some ideas about their ID’s, but am not completely sure. Can anyone help me with their identification?
This first anole was found on a tree trunk near the stream at night, while the second one was found on a pole in the morning. Both Anolis lionotus?
This third one was found on a low shrub in the morning. Anolis limifrons?
The fourth anole was found on a low-growing shrub at night. Anolis lemurinus?
Shane Campbell-Staton had fortuitously measured the thermal physiology of a number of populations of the green anole, Anolis carolinensis, the summer before 2014’s Polar Vortex. So, he went back and examined the survivors. And sure enough, in the most southerly populations, those most strongly affected by the cold snap, natural selection had occurred. Shane tells Scientific American all about it in this podcast. The nifty figure above comes from the University of Illinois’ press release.
We reported recently that knight anoles (Anolis equestris) have shown up in the T&C. Here’s more on the story from B Naqqi Manco, the Terrestrial Ecologist at the Department of Environment and Maritime Affairs, Turks and Caicos Islands Government:
Cuban knight anoles are currently known from two sites on Providenciales: Vicinity of Beaches Resort in The Bight and Amanyara Resort on Northwest Point. Both populations showed up after the importation of large trees for landscaping from Miami. The properties are both irrigated pretty heavily to keep the bigger trees going. The tree imports were brought in before the Department of Agriculture was fully operative, so unfortunately things got in at that time that probably shouldn’t have made it through.
I don’t have confirmation of the knight anoles breeding, but I know The Bight population has been spreading with individuals having been found on adjacent properties and in a nearby residential neighbourhood. I would be very surprised if they’re not breeding on either site. Unfortunately we don’t have the capacity to monitor them well but this is something we want to keep a closer eye on and it would make a worthwhile research project for a student or intern.
Thus far, they have not been reported from any other island or cay.
The paper, by Nancy Staub and Rachel Mueller and just out in Copeia, is a delightful biography of DBW, as he is referred to by his lab and many others. As for the anole bit, you’ll have to read it to find out.
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.
Travis in the Dominican Republic with Anolis fowleri. Photo by Luke Mahler.
Two recent talks at JMIH 2017 shed light on key morphological characters in anoles: toe pad shape and limb length. Travis Hagey presented his work which looks to shed light on why lizard toe pads are shaped the way that they are and addresses whether gecko and anole toe pads are convergent structures. Working with a team of undergraduates, Travis used geometric morphometrics to analyze the structure of toepads in a diverse group of geckos and anoles. Travis found that anole and gecko toe pads have a similar range of values for traits such as the placement of pads on the toes and the shape of the toes (skinny or fat) in relation to claws. However, anole toe pads formed a distinct cluster indicating that they occupy a unique area of trait space not used by geckos. This finding suggests that the divergent evolutionary history of anoles and geckos has resulted in independent evolutionary explorations of toe pad shape.
Immediately following Travis’ talk, Robin Andrews presented work investigating the embryological development of morphological characters in diverse lizard species. In anoles, consistent differences in the morphology of divergent species support the existence of different anole ecomorphs. Previous research by Sanger and colleagues has shown that the differences in limb-length between anoles of different ecomorphs have their origins early in embryonic development. These early differences in limb length continue throughout the development of anoles into hatchlings and adult forms, a pattern known as transpositional allometry.
Robin compared patterns of limb, tail, and head growth in early stage embryos of four different lizard species, including a chameleon, two geckos, and the brown anole (Anolis sagrei). She found that species-specific differences in limb and tail lengths were exhibited as soon as limb and tail buds emerged from the body and were both best characterized by the same pattern, transpositional allometry. Embryonic head growth, however, showed no specific pattern. Robin’s findings suggest that the adaptive evolution of adult morphology in anole ecomorphs as well as other diverse lizard species is underpinned by developmental reprogramming.
Travis Hagey, Jordan Garcia, Oacia Fair, Nikki Cavalieri, and Barb Lundrigan: Variation in Lizard Adhesive Toe Pad Shape Robin Andrews: Developmental Origin of Limb Size Variation in Lizards
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.