We’ve seen anole wedding cakes and thesis defense cakes, but here’s a new one. Anole research veteran Natalie Jacewicz reports:
For my bachelorette party, my bridesmaids went to an erotic bakery (quite the business niche) in Boston and brought the shop pictures of Anolis lizards. The bakery evidently usually deals in, er, human encounters, so only had skin-toned frosting, and the store clerks weren’t sure if they could do anything lizard themed. But the shop owner evidently got really into the project, did a lot of independent anole research, and produced the cake below. Yes, that is a bridal veil on the yellow one.
Some weeks ago, a paper I wrote on the display behaviour and morphology of fan-throated lizards was published early online at the Journal of Herpetology. Some unfortunate timing meant that my paper did not incorporate these lizards’ new taxonomy, recently published by V. Deepak and colleagues. In this post, I’m going to summarize my results, and explore them in the context of what we now know about Sitana (Agamidae) systematics.
Male fan-throated lizards (surprise, surprise) have fans under their throats that are displayed in a manner analogous to the Anolis dewlap. The appearance of the throat-fan varies dramatically across this group, from small and mostly white to large and blue, black, and orange. I wanted to answer two broad questions
- Does display behaviour vary with throat-fan morphology? In other words, if you have different tools with which to communicate, do you communicate differently?
- Can we examine morphological and environmental variation to deduce anything about how this variation in throat-fan morphology has evolved?
To address these two questions, I measured the display behaviour, morphology, and environment of eight populations of lizards, from three “throat-fan variants.” I found the following:
- The main axis of variation in display behaviour differed between the coloured-fan variant and everybody else. Displays were fewer and longer in the coloured-fan variant, and included more head twists. The same axis of display behaviour did not differ between the white-fan and the intermediate-fan variants, though there was variation in the frequency of head-bobs across populations with different-sized throat-fans. These differences in display behaviour make sense in light of morphology. Head twisting was more frequent in the variant with a large blue section on the throat-fan that appears iridescent. Head-bobs, which often co-occur with a fully extended throat-fan, were more frequent in the variant(s) with smaller throat-fans (see Figure 6 in my paper for more).
- Throat-fan elaboration (both size and colour) was paired with increased male-biased sexual size dimorphism, suggesting sexual selection as a likely selective force driving throat-fan variation.
- Habitat structure did not co-vary with throat-fan morphology, suggesting that the visual environment is unlikely to play much of a role in the maintenance of this variation in throat-fan morphology. But because these lizards all persist in human-modified landscapes, it is difficult to discern how important the visual environment was for the origin of dewlap diversification in this group.
Based on geography, I can tell that all three of the coloured-fan variant populations I sampled belong to the newly described Sarada darwinii. The white-fan populations are Sitana laticeps and Sitana spinaecephalus (+ one population I’m not sure about), and the northern and southern intermediate-fan populations are Sitana ponticeriana and Sitana visiri respectively. Recast in terms of these species delimitations, I found that:
- Display behaviour differs between the genera Sitana and Sarada. It doesn’t vary consistently with species within Sitana, though variation in head-bobbing should be explored further.
- There are two broad possibilities for throat-fan evolution in the group. One possibility is that throat-fan elaboration and a shift towards male-biased SSD has evolved independently twice, once in Sarada (Clade 1) and once in the South India/Sri Lanka clade (Clade 3 in the phylogeny) in Sitana. The other possibility is the reduction of dewlap size and colour in the west Indian Sitana clade (Clade 2). This question won’t be definitively answerable until we have a phylogeny that includes the remaining north-eastern species of Sitana as well as more species of the sister genus Otocryptis, which also vary in the presence and morphology of the throat-fan.
Before knowing about the phylogeny, I predicted that throat-fan elaboration had evolved twice in fan-throated lizards, based on a suite of differences between the coloured-fan variant (now Sarada) and the intermediate-fan variant (now Sitana Clade 3). The main ones are:
- Different display behaviour.
- Different allometric relationships between body size and throat-fan size, suggesting different ways in which throat-fans have gotten big.
- Different spectral reflectances from the blue and orange patches, plus the presence/absence of black on the throat-fan.
- The ability of Sitana, but not Sarada, to turn “on” and “off” the blue colour on their throat-fans (more about this in a future post!).
These differences now lead me to favour the first of the two possibilities outlined above: repeated, somewhat parallel evolution of throat-fan elaboration, as opposed to the loss of an elaborate throat-fan. Given that the sister genus Otocryptis has also either evolved or lost a throat-fan (throat-fans are present in O. nigristima and O. wiegmanni but not O. beddomi), this group is positively rife with lability in display evolution, offering all sorts of exciting possibilities for future research!
Over on phostracks.com: Florida Wildlife, Ecology and More, Janson Jones reports on a very successful trip to Miami that yielded many anoles, most notably the Jamaican crown-giant, Anolis garmani and the Cuban knight anole, Anolis equestris.
Janson Jones is at it again. Actually, he’s been at it for a year, but somehow that slipped below our radar. The former purveyor of Dust Tracks on the Web has a new venue, phosTracks.com: florida wildlife, ecology and more.
Like it’s predecessor, phosTracks is full of keen natural history, engagingly presented and complemented by gorgeous photography. And better yet, anoles are one of Jones’ two favorite animals, neck-and-neck (hard as it may be to believe) with watersnakes.
Check out some of Jones’ recent musings on:
and more! Stay on these pages for some of his giant anole goodness coming up soon!
At JMIH 2016, I chatted with Johanna Wegener, a graduate student at the University of Rhode Island in Jason Kolbe’s lab, about her poster detailing her work identifying hybridization between Anolis carolinensis and A. porcatus in southern Florida.
Interspecific hybridization in anoles is thought to be fairly rare, with the best-known example being hybridization between Anolis carolinensis (native to the southeastern U.S.) and A. porcatus (native to Cuba) in southern Florida. I was surprised to learn how little we know about this rumored hybrid zone.
A. porcatus was likely introduced into Florida within the last few decades, but the striking morphological similarities between A. carolinesis and A. porcatus make anecdotal reports of hybridization hard to confirm. Wegener conducted the first genetic analyses of hybridization between A. carolinesis and A. porcatus. She genotyped 18 nuclear microsatellites from green anoles in Florida (Palm Beach and South Miami) and western Cuba and conducted a STRUCTURE analysis and found support for three genetic clusters consisting of Cuban A. porcatus, and two Floridian groups (one from Palm Beach and one from South Miami). With the addition of the mitochondrial ND2 marker, she found that the South Miami population had both A. carolinensis and A. porcatus haplotypes. Interestingly, there appeared to be very few recent hybrids; instead, the hybrid group appeared distinct from either parent group, suggesting that hybridization has been occurring for several generations.
In addition, Wegener looked at the variation in A. porcatus and A. carolinensis markers in each hybrid individual and found examples of some parent markers being retained at high proportions in the hybrids, possibly suggesting the retention of beneficial parent alleles in the hybrids.
Given that this study was only conducted at two sites in Florida, the exciting next step of this study is to better quantify the genetic makeup of hybrids across southern Florida and map out the hybrid zone.
De Gao and Gad Perry have recently detected the small island effect (SIE) and nestedness patterns of Anolis Lizards of the West Indies. We applied regression-based analyses, including linear regression and piecewise regressions with two (two-slope function and left-horizontal with one threshold function) and three (three-slope function and left-horizontal with two thresholds function) segments, to detect the SIE and then used the Akaike’s information criterion (AIC) as a criterion to select the best model. We used the NODF (a nestedness metric based on overlap and decreasing fill) to quantify nestedness and employed two null models to determine significance. Moreover, a random sampling effort was made to infer about the degree of nestedness at portions of the entire community.
We found piecewise regression with three segments performed best, suggesting the species–area relationships (SARs) possess three different patterns that resulted from two area thresholds: a first one, delimiting the SIE, and a second one, delimiting evolutionary processes. Moreover, the traditional two-segment piecewise regression method may cause poor estimations for both slope and threshold value of the SIE. Thereby, we suggest previous SIE detection works that conducted by two-segment piecewise regression method, ignoring the possibility of three segments, need to be reanalyzed. Anti-nestedness occurred in the entire system, whereas high degree of nestedness could still occur in portions within the region. So, nestedness may still be applicable to conservation planning at portions even if it is anti-nested at the regional scale.
I was recently in Abaco, Bahamas with Losos lab post-doc Oriol LaPiedra and Ph.D. candidate Darío Fernández-Bellon from University College Cork, Ireland, to carry out some behavioral studies of Anolis sagrei on the island and its surrounding small cays. We kayaked (a highly recommended transportation mean for its lesser-impact on the marine ecosystem, not having to rely on the tide schedule, while allowing you to see rays and sharks and sea turtles!) our way out to islands that are known to have A. sagrei naturally existing alone, or with one of their natural predators, Leiocephalus carinatus.
Curly-tailed lizards are known to prey on A. sagrei and can have significant impact on anole behavior and adaptation. Twice I observed Leiocephalus capturing and consuming A. sagrei, one of which was an adult male and the other an adult female. We have also noticed that the A. sagrei on these island tend to perch higher and are seldomly seen on rocks or leveled ground compared to those on islands without curly tails, so this behavior could be an effect of Leiocephalus being present.
On a different island where Leiocephalus were absent, A. sagrei are still under predation pressure, this time by red-winged blackbirds nesting on the island. We observed a female blackbird with an A. sagrei in its beak waiting for us to leave the island so that it can feed its chicks. This observation suggests that A. sagrei on islands without Leiocephalus might still be under predation pressure by other species that might not be present on the island at all times. Also, predation pressure exerted by an aerial predator differs from that by a terrestrial predator or if both predators are present, so this might be a factor in morphological or behavioral changes in these lizards on these islands.
Other interesting observations include A. sagrei density on islands seems to be unintuitive. Some small islands with fewer perches hosted many more adult males and females than large islands did. Sizes of individuals also seem to vary greatly between different islands: small cay A. sagrei seem to be, on average, larger than those on mainland Abaco. Personally, I am unable to note major differences between islands which might have resulted in these observations. I’m excited to see if the data we’ve collected will give more insight into these observations as well as other behavioral results that will come from this study!
Anole stalwart Greg Mayer gave a wonderful talk discussing the distribution and morphology of the large and maybe-extinct Anolis roosevelti. A. roosevelti, commonly known as the Culebra Island giant anole, was first described in 1931 by Chapman Grant, a US Army Major and practising herpetologist, from a single adult male specimen collected on Culebra. Although Reinhardt and Lutken, in 1863, had already provided an accurate description of A. roosevelti, but under an alternative name of A. velifer.
Reinhardt and Lutken’s specimens were collected from Vieques, Tortola, and St. John, although Greg having the opportunity to study them meant tracking them down to natural history collections in both Copenhagen and Stockholm. In total, this entire species is known from eight specimens, only six of which are still in existence (Greg had the opportunity to study all six, meaning he’s now seen more roosevelti than any other anolologist?). Greg explains that roosevelti based on the limited information provided by Dimas Villanueva, who collected the holotype, and his own investigations, roosevelti can be classified as a “crown-giant” ecomorph. This means that the eastern islands of the Puerto Rico bank had a series of four ecomorphs, with roosevelti being what Ernest Williams termed a climatic vicariant of cuvieri, occuring in (and presumably being adapted to) the more xerophytic forests of the eastern bank islands.
Greg went on to describe the morphological features which distinguish A. roosevelti from a A. cuivieri, an ecologically and morphologically similar species from neighbouring Puerto Rico. Roosevelti is a larger, brownish gray rather than green as is seen in cuvieri (although check out these gray cuvieri preveiously mentioned on AA). Roosevelti generally has larger head scales, and a more elongate and deeply grooved head – these differences are confirmed in the ANCOVA analyses below.
So, what chances are there of seeing roosevelti in the wild? Low, probably. No specimens have been collected since 1932, and several researchers, including Greg, have recently scoured both Vieques, St. John and Tortola but with no success. By far the most extensive searches have been conducted by Ava Gaa, who exhaustively searched Culebra (totalling 1500 hours of looking!) as well as short visits to Vieques and St. John all with no success. Tantalising reports of potential candidates turned out to be juvenile green iguanas. Greg concludes by recommending that the long-protected and relatively poorly explored eastern half of Vieques may hold the secret to if any populations remain.
Travis Hagey presented some new results from his ongoing research on the evolution of functional traits in lizards. Travis normally works on geckos, but frequently includes Anolis species in his studies. Last year at Evolution, Travis told us about toepad evolution by comparing gecko toepads to those of anoles and skinks. Along the same vein, this year at JMIH Travis talked about patterns of limb-length across different lizard groups.
Travis started with anoles as an example of morphology being correlated with habitat use. As we all know, anole limb length is associated with structural habitat. Lizards like Anolis occultus (a twig anole) use thin perches and have very short legs. Other species that perch on broader substrates tend to have longer legs. Travis is interested in finding out if this pattern holds for other groups of lizards.
He started by comparing anoles to geckos to see if relative limb length differed between the groups. He accumulated an impressive database of hindlimb lengths from many gecko and anole species and when he looked at the relationship between hindlimb length and body size (SVL), he found that for a given body size anoles tended to have longer limbs than equivalently sized geckos. He then added in data for a number of species from Liolaemus, Tropidurus, and Phrynosomatidae. Interestingly, he found that these other groups all clustered with the anoles. This suggests that there are possibly two relationships between limb-length and body size across lizards.
Travis ended by commenting on how this might relate to habitat use. He analyzed hindlimb length by perch diameter for anoles (red line) and geckos (black line). Geckos, it turns out, have a different relationship between perch use and limb length than anoles: geckos with shorter limbs tend to use broader diameter perches! Travis is still working on this research and is looking for data on limb length for many groups. If you have hindlimb length data from lizards you should email Travis to help out!